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EDITORIAL ADVISORY BOARD
PETER BUCHANAN (2011-2017), Chair
Auckland, New Zealand
SABINE HUHNDORE (2011-2016), Past Chair
Chicago, Illinois, U.S.A.
BRANDON MATHENY (2013-2018)
Knoxville, Tennessee, U.S.A.
KAREN HANSEN (2014-2019)
Stockholm, Sweden
ISSN 0093-4666 (PRINT)
ISSN 2154-8889 (ONLINE)
MYCOTAXON
THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY & NOMENCLATURE
OCTOBER-DECEMBER 2017
VOLUME 132 (4)
http://dx.doi.org/10.5248/132-4
EDITOR-IN-CHIEF
LORELEI L. NORVELL
editor@mycotaxon.com
Pacific Northwest Mycology Service
6720 NW Skyline Boulevard
Portland, Oregon 97229-1309 USA
NOMENCLATURE EDITOR
SHAUN R. PENNYCOOK
PennycookS@LandcareResearch.co.nz
Manaaki Whenua Landcare Research
Auckland, New Zealand
MyYcoTAxONn, LTD. © 2017
www.mycotaxon.com &
www.ingentaconnect.com/content/mtax/mt
P.O. BOX 264, ITHACA, NY 14581-0264, USA
IV ... MYCOTAXON 132(4)
MYCOTAXON
VOLUME ONE HUNDRED THIRTY-TWO (4) — TABLE OF CONTENTS
132-4: TABLE OF CONTENTS, NOMENCLATURAL UPDATES, PEERS, EDITORIAL
Nomenclatural novelties & typifications ........ 00. c eee eee vii
PREV ICWALS nc cc EOS to cae Sar oto. wee et vate es Re Ae hettele «gates ix
ERPACE UL PPOVIOUS ASUS. SE Ae an at Ls ae ee hy eh ae hd ee ee x
PROVDEEEAIEOV: 30.4 usariga cre tee iscrinh ence tte nry: ey E ty Aas CBO Th aoe cd ecarern gg 8 a xi
ZOES AU UPAISSIOMPKO CCAS ng Sse tis, auiklace be ayd chi ine Pach iechtace title SY xiii
RESEARCH ARTICLES
Russula vinosoflavescens sp. nov.,
from deciduous forests of Northern Alsace, France
JEAN MICHEL TRENDEL, FELIX HAMPE & ANNEMIEKE VERBEKEN 707
Triadelphia acericola and T: centroseptata spp. nov.,
and a synopsis of the genus DeE-WE! LI & JIAN-REN YE 723
Glischroderma Fuckel Greécorre L. HENNEBERT 745
Ellismarsporium gen. nov. and Stanhughesiella gen. nov. to
accommodate atypical Helminthosporium and Corynesporella species
RAFAEL FE, CASTANEDA-Ru1IZ, DE-WEI LI, XIU-GUO ZHANG, BRYCE KENDRICK,
BEATRIZ RAMOS-GARCIA, SIMON PEREZ-MARTINEZ & DAYNET Sosa 759
Endophragmiella jiulingensis sp. nov.
and two new records from southern China Hao-Hva Lt, Kai ZHANG,
CHUN-LING YANG, JI-WEN XIA & XIU-GUO ZHANG 767
Pseudocercospora lysidices sp. nov. on Lysidice rhodostegia
from China FENGYAN ZHAI, YINGLAN Guo, YINGJIE Liu & HONGLIAN Li 773
Bactrodesmiastrum domesticum sp. nov. and
Conioscypha varia from indoor environments
DE-WEI LI, CHIN S. YANG & ARIUNAA JALSRAI 779
Russula brunneovinacea sp. nov.,
from northeastern China Xu-MENG JIANG, YANG-Kun L1,
JUN-FENG LIANG & JIAN-RONG Wu 789
Amanita pallidorosea in Pakistan and
its ectomycorrhizal association with Quercus oblongata
MUNAZZA KIRAN, JUNAID KHAN, AROO] NASEER,
HASSAN SHER & ABDUL NasiIR KHALID 799
Ellisembia pseudokaradkensis sp. nov.
from Hainan, China Min Qiao, XING Du,
ZHAO-HuI BIAN, JIE PENG & ZE-FEN YU 813
OCTOBER-DECEMBER 2017... V
Colpoma guadueticola sp. nov. in a guadual forest
from Quindio Department, Colombia TANIA RAYMUNDO,
RICARDO SOTO-AGUDELO & RICARDO VALENZUELA 819
Baeomyces lotiformis sp. nov. from China
SHUNAN CAO, JIANFENG HE, FANG ZHANG, HUIMIN TIAN,
CHUANPENG Liv, HAIYING WANG & QIMING ZHOU 831
Lirula sichuanensis sp. nov. on Picea likiangensis var. rubescens
from Sichuan, China
CuHuUN-LIN YANG, XIU-LAN Xu, ZHENG-GAO ZHANG & YING-GAO Liu 839
Hypoderma rubi on two new hosts in Slovakia
MARTIN PasTIRCAK & KATARINA PASTIRCAKOVA 849
New reports of Myriospora (Acarosporaceae) from Europe
KERRY KNUDSEN, JANA KOCOURKOVA & ULF SCHIEFELBEIN 857
Perenniporia puerensis sp. nov. from southern China Wet-Li Liv,
TaI-MIN Xu, SHAN SHEN, XIANG-Fu Liu, YANG SUN & CHANG-LIN ZHAO 867
Sporidesmiopsis lushanensis sp. nov. from Lushan Mountain, China
X1A0-MEI WANG, ZI-JIAN ZHAO, SHAN-SHAN CHEN,
X1A0-MaAN Li, Hao-Hua LI, X1u-Guo ZHANG & JI-WEN X1A 875
Repetophragma elegans sp. nov. from Hainan Province, China
X1AO-MEI WANG, SHAN-SHAN CHEN, XIAO-MAN LIv,
ZI-JIAN ZHAO, Hao-Hua LI, X1u-GUO ZHANG & JI-WEN XIA 881
A contribution to the study of Helotiales and Rhytismatales in Turkey
MAKBULE ERDOGDU, GOKHAN DoGaN, EL$AD HUSEYIN & ZEKIYE SULUDERE 885
Huneckia pollinii and Flavoplaca oasis newly recorded from China
CoNG-ConG MIA, XIANG- XIANG ZHAO,
ZUN-TIAN ZHAO, HURNISA SHAHIDIN & Lu-Lu ZHANG 895
Paliphora bicolorata sp. nov. from the Brazilian Atlantic Forest
ELAINE MALosso, PHELIPE M.O. Costa,
MarcE La A. BARBOSA, GABRIELA V.R. Da SILVA & RAFAEL E. CAsSTANEDA-Ru1z 903
Inocybe shawarensis sp. nov. in the Inosperma clade from Pakistan
A. NASEER, A.N. KHALID & MATTHEW E. Smi1TH 909
Phaeomonilia aquatica sp. nov.,
an aquatic hyphomycete from China Jun-EN Huana, Hat-Yan Sona,
X1-GEN HUANG, JIAN Ma & DiAn-Minc Hu 919
Minimelanolocus atlanticus sp. nov. and M. navicularis
from the Brazilian Atlantic Forest | PHELIpE M.O. Costa, ELAINE MALosso,
MarceELa A. BARBOSA, WANDERSON L. TAVARES & RAFAEL FE. CASTANEDA-RuIz 925
New records of Amanita
from Tehuacan-Cuicatlan Biosphere Reserve, Mexico
EVANGELINA PEREZ-SILVA & ABRAHAM J. MEDINA-ORTIiz 933
vi ... MYCOTAXON 132(4)
Thelephora iqbalii sp. nov.
from the Himalayan moist temperate forests of Pakistan
ABDUL NASIR KHALID & MUHAMMAD HaAnlir 943
Three new combinations in Drepanopeziza
for species on poplar Amy Y. RossMAN, W. CAvVAN ALLEN,
Lisa A. CASTLEBURY & GERARD VERKLEY 951
Key to the lichen families Pyrenulaceae and Trypetheliaceae
in Vietnam, with eight new records
SANTOSH JOSHI, D.K. UPRETI & JAE-SEOUN Hur 957
Acumispora delicata sp. nov.
from the Brazilian Atlantic Forest
PHELIPE M.O. Costa, MARCELA A. BARBOSA, WANDERSON L. TAVARES,
DAYNET SOSA, SIMON PEREZ-MARTINEZ, RAFAEL FE. CASTANEDA-RUIZ
& ELAINE MALosso 971
Anisogenispora insignissima gen. & sp. nov.
from the Brazilian semi-arid region = SHEILA MIRANDA LEAO-FERREIRA,
Luis FERNANDO PASCHOLATI GUSMAO & RAFAEL F. CASTANEDA-RUIZ 977
REGIONAL MYCOBIOTAS NEW TO THE MYCOTAXON WEBSITE 985
Diversity of wood-inhabiting aphyllophoraceous basidiomycetes
on the island of Cyprus MICHAEL LOIZzIDES
Checklist of saprobic asexual microfungi from the
tropical montane cloud forest of Veracruz, México
Rosa Maria ARIAS, GABRIELA HEREDIA & RAFAEL F. CASTANEDA-RUIZ
BOOK REVIEWS AND NOTICES LORELEI NORVELL & ELSE VELLINGA (zps.) 987
AGARICUS OF NORTH AMERICA
(Richard W. Kerrigan; 2016)
HEBELOMA (FR.) P. KUMM.
(H.J. Beker, U. Eberhardt & J. Vesterholtt; 2016)
LICHENS OF MEXICO: THE PARMELIACEAE
(Maria Herrera-Campos, Rosa Emilia Pérez-Pérez, Thomas H. Nash II], eds.; 2016)
PUBLICATION DATE FOR VOLUME ONE HUNDRED THIRTY-TWO (3)
MYCOTAXON for JULY-SEPTEMBER 2017, (I-xIv + 471-706)
was issued on October 2, 2017
OCTOBER-DECEMBER 2017...
NOMENCLATURAL NOVELTIES AND TYPIFICATIONS
PROPOSED IN MYCOTAXON 132(4)
Acumispora delicata P.M.O. Costa, Malosso & R.E Castaneda
[IF 554059], p. 972
Anisogenispora S.M. Leao, Gusmao & R.F. Castaneda
[IF 552545], p. 978
Anisogenispora insignissima S.M. Leao, Gusmao & R.F. Castafeda
[IF 552546], p. 978
Bactrodesmiastrum domesticum D.W. Li & Chin S. Yang
[MB 820855], p. 780
Baeomyces lotiformis S.N. Cao
[MB 819009], p. 834
Colpoma guadueticola Raymundo, Soto-Agudelo & R. Valenz.
[MB 820664], p. 822
Drepanopeziza brunnea (Ellis & Everh.) Rossman & W.C. Allen
[MB 822026] (lectotypified), p. 952
Drepanopeziza castagnei (Desm. & Mont.) Rossman & W.C. Allen
[MB 822027] (lectotypified), p. 953
Drepanopeziza populi (Lib.) Rossman & W.C. Allen
[MB 822028] (lectotypified), p. 954
Ellisembia pseudokaradkensis M. Qiao & Z.E. Yu
[MB 820988], p. 814
Ellismarsporium R.F. Castahtleda & X.G. Zhang
[MB 807603], p. 760
Ellismarsporium catenatum (Matsush.) R.F. Castafieda & X.G. Zhang
[MB 807605], p. 763
Ellismarsporium hypselodelphyos (M.B. Ellis) R.E Castaheda & X.G. Zhang
[MB 807604], p. 763
Ellismarsporium parvum R.F. Castaneda & W. B. Kendr.
[MB 807606], p. 763
= Helminthosporium parvum R.F. Castaneda & W.B. Kendr.,
nom. illeg. (non Grove 1886)
Ellismarsporium pinarense (R.F. Castaneda) R.E. Castaneda & X.G. Zhang
[MB 807607], p. 763
Ellismarsporium senseletii (Bhat & B. Sutton) R.E Castafeda & X.G. Zhang
[MB 807608], p. 763
Ellismarsporium simpliphorum (Matsush.) R.F. Castafieda & X.G. Zhang
[MB 807609], p. 763
Ellismarsporium zombaense (B. Sutton) R.F. Castafeda & X.G. Zhang
[MB 807610], p. 763
VII
vill ... MYCOTAXON 132(4)
Endophragmiella jiulingensis H.H. Li & X.G. Zhang
[MB 823127], p. 769
Inocybe shawarensis Naseer & Khalid
[MB 820130], p. 912
Lirula sichuanensis X.L. Xu, C.L. Yang & Y.G. Liu
[MB 819592], p. 841
Minimelanolocus atlanticus P.M.O. Costa, Malosso & R.F. Castaneda
[IF 554056], p. 926
Paliphora bicolorata Malosso, P.M.O. Costa, & R.FE. Castaneda
[MB 823671], p. 904
Perenniporia puerensis C.L. Zhao
[MB 823635], p. 868
Phaeomonilia aquatica J.E. Huang, H.Y. Song, Jian Ma & D.M. Hu
[MB 821142], p. 920
Pseudocercospora lysidices Y.L. Guo & F.-Y. Zhai
[MB 819544], p. 776
Repetophragma elegans J.W. Xia & X.G. Zhang
[MB 821043], p. 883
Russula brunneovinacea X.M. Jiang, Yang K. Li & J.F. Liang
[MB 819154], p. 791
Russula vinosoflavescens Trendel & F. Hampe
[MB 819428], p. 711
Sporidesmiopsis lushanensis J.W. Xia & X.G. Zhang
[MB 823681], p. 876
Stanhughesiella R.F. Castaieda & D.W. Li
[MB 807614], p. 764
Stanhughesiella dictyoseptata (S. Hughes) R.F. Castaneda & D.W. Li
[MB 807615], p. 764
Thelephora iqbalii Khalid & Hanif
[MB 800830], p. 945
Triadelphia acericola D.W. Li
[MB 813008], p. 725
Triadelphia archontophoenicicola D.W. Li
[MB 813055], p. 728
= Triadelphia australiensis Joanne E. Taylor, K.D. Hyde & E.B.G. Jones
2003 nom. illeg. non B. Sutton 1989)
Triadelphia centroseptata Venkateshw., S.M. Reddy & S.R. Reddy ex D.W. Li
[MB 823119], p. 731
= Triadelphia centroseptata Venkateshw., S.M. Reddy & S.R. Reddy,
1997 [“1996”], nom. inval.
OCTOBER-DECEMBER 2017...
REVIEWERS — VOLUME ONE HUNDRED THIRTY-TWO (4)
The Editors express their appreciation to the following individuals who have,
prior to acceptance for publication, reviewed one or more of the papers
prepared for this quarter.
Najam ul Sehar Afshan
André Aptroot
Juliano M. Baltazar
Annarosa Bernicchia
Uwe Braun
Rafael F. Castahieda —Ruiz
Bao-Kai Cui
Gregorio Delgado
Ursula Eberhardt
Martin Esqueda
Mark L. Gleason
Shouyu Guo
Luis Fernando Pascholati Gusmao
Rosanne Healy
Gabriela Heredia Abarca
Kathie T. Hodge
Sana Jabeen
Peter R. Johnston
Jason Karakehian
Geoffrey Kibby
Sevda Kirbag
S. Kondratyuk
James C. Lendemer
De-Wei Li
Yan-Chun Li
Ying-Ren Lin
Robert Lticking
Jian Ma
P. Brandon Matheny
John W. McCarthy
Eric H.C. McKenzie
David W. Minter
Josiane S. Monteiro
Gabriel Moreno
Abdul Rehman Niazi
Lorelei L. Norvell
Shaun R. Pennycook
Sergio Pérez Gorjon
Elias Polemis
Hyeon-Dong Shin
H.J.M. Sipman
Jeffrey Stone
Else C. Vellinga
Andrew W. Wilson
Eugene Yurchenko
Huang Zhang
Xiu-Guo Zhang
Huan-Di Zheng
X ... MYCOTAXON 132(4)
ERRATA FROM PREVIOUS VOLUMES
VOLUME 122
pp. 449-460 For: Arthrinium rasikravindrii or A. rasikravindrii
READ: Arthrinium rasikravindrae or A. rasikravindrae
The paper “Arthrinium rasikravindrii [sic] sp. nov. from Svalbard, Norway” by Singh et al.
proposed a new species honoring Dr. Rasik Ravindra. Although the director of the National
Centre for Antarctic & Ocean Research and leader of the first Indian Arctic Expedition is
a man (as indicated by the masculine genitive ending -ii on the published species epithet),
the INTERNATIONAL CODE OF NOMENCLATURE FOR ALGAE, FUNGI, AND PLANTS requires the
feminine genitive ending —ae for any name ending in ~a, regardless of the gender of the person
(Rec. 60C.1(a)).
The misspelled species epithet should be corrected to ‘rasikravindrae’ throughout. The
misspelling appears on p. 449 lines 5, 13, 19; in running titles on pp. 451, 453, 455, 457, 459; on
p. 451 line 5; p. 452 line 13; p. 453 Fie. 1 legend, line 1; p. 454 lines 31, 33, 36 and Fia. 2 legend,
line 2; p. 456, Phylotree (Fic. 3): top 7 branches; Fie. 3 legend, line 2; primary text line 5; p. 457
Phylotree (Fic. 4): branches 22-28; p. 458 lines 2, 7, 9, 10, 12, 14, 17 & 18.
VOLUME 132(3)
p. 589 line 3 App: Mycoraxon, Ltd. is thanked for underwriting the publication costs
for this manuscript.
Back cover, line 1
FOR: Cephalotrichum tuffiforme READ: Cephalotrichum turriforme
OCTOBER-DECEMBER 2017 ... XI
FROM THE EDITOR-IN-CHIEF
MATERIALS & METHODS— REMOVING THE TEDIUM. Mycologists and other scientists are
usually introduced to materials and methods while writing lab reports, dissertations,
or grant applications—situations in which devotion to minute detail is encouraged
or where a budget must be calculated. When publishing short papers, however, the
‘Me&w’ section is best kept brisk, pertinent, and succinct. Only procedures producing
results should be included. Although researchers may have spent many, many
exasperating hours extracting, amplifying, and sequencing DNA, no one wishes to
read about the process unless phylogenetic results appear later in the paper.
Likewise, if the procedures followed are the same found in a previous publication,
do not invent text or copy from the older paper; simply insert a text reference, such
as “ITS rDNA was extracted and amplified according to White & al. (1990).” Once.
Extra (brief) detail is needed only when the referenced protocol has been modified.
Easy worldwide access to today’s Internet means that Mycotaxon readers are no longer
heart-warmed by recitation of company, city, country from which a particular piece lab
equipment was acquired. Authors are encouraged to place Brand and Model number
(no ® symbol required!) before the equipment. Thus, microscopical measurements
should be taken using NOT “..a compound light microscope (Leica®, Model DM RB,
Leica Microsystems (Danaher), Wetzlar, Germany) loaded with Nomarski optics...” but
simply “...a Leica DM RB compound microscope with DIC optics.”
And please, people! In this day of chemical, cultural, molecular, and population
analyses, there is no longer a need to encompass “morphological” and “anatomical”
using the needlessly long term: “morpho-anatomical.”. “Morphology” which covers
both anatomy as well as features visible to the naked eye, may be used all by itself!
[Species circumscribed relying only on anatomical characters are still understood to be
based on a ‘morphological species’ concept.]
DIAMETER VS. WIDTH—We know. It is extremely tempting to refer to the short
dimension of a hypha measured with 1000x-power as a ‘width (which contrasts with
the long dimension, the ‘length’). Unfortunately using ‘width’ for diameter conveys
the impression that the object measured is two-dimensional. It isn’t—it is three-
dimensional, and what you have measured is the diameter. Rather than write out ‘in
diameter’ in your technical descriptions and keys, try placing diam. (or even diam
without the full stop) after the unit (e.g.: “23 um diam” or “23 um diam.”). MycoTAxoNn
accepts both styles but does request consistency! This has the added bonus of reminding
readers that what we see through a microscope lens only sEEMs to exist in one plane.
MYCOTAXON 132(4) presents 31 research papers by 132 authors representing 18
countries and revised by 48 expert reviewers, adds two new mycobiotas to its
website, and reviews three excellent books published in late 2016.
XII ... MYCOTAXON 132(4)
Within its pages are THREE new genera (Anisogenispora, Ellismarsporium,
Stanhughesiella) and 22 species new to science representing Acumispora, Anisogenispora,
Minimelanolocus, and Paliphora from Brazil; Bactrodesmiastrum and Triadelphia
from the United States; Baeomyces, Ellisembia, Endophragmiella, Lirula, Perenniporia,
Phaeomonilia, Pseudocercospora, Repetophragma, Russula, and Sporidesmiopsis from
China; Colpoma from Colombia; Inocybe and Thelephora from Pakistan; and Russula
from France.
We also offer 12 new combinations or replacement names in Drepanopeziza,
Ellismarsporium, Stanhughesiella, and Triadelphia and keys to species in Acumispora,
Ellismarsporium, Phaeomonilia, Pyrenulaceae & Trypetheliaceae in Vietnam,
Sporidesmiopsis, and Triadelphia.
Range extensions and/or new hosts for previously named taxa are cited for Myriospora
in the Czech Republic & Italy; Helotiales & Rhytismatales in Turkey; Huneckia &
Flavoplaca in China; and Amanita in Mexico’s Tehuacan-Cuicatlan Biosphere Reserve.
Phylogenetic support and the ectomycorrhizal association with Quercus are confirmed
for Amanita pallidorosea in Pakistan while Hypoderma rubi is shown on two new hosts
in Slovakia.
Finally, in his clarification of Glischroderma, Mycotaxon co-founder Hennebert
untangles a long-standing confusion within Glischroderma-Pachyphlodes-Ostraderma
in honor of his former collegue and fellow co-founder, Dick Korf.
Warm regards,
Lorelei L. Norvell (Editor-in-Chief)
10 January 2018
OCTOBER-—DECEMBER 2017... XIII
2018 MYCOTAXON SUBMISSION PROCEDURE
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Mycotaxon publishes four quarterly issues per year. Both open access and
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 707-721
https://doi.org/10.5248/132.707
Russula vinosoflavescens sp. nov.,
from deciduous forests of Northern Alsace, France
JEAN MICHEL TRENDEL’, FELIX HAMPE? & ANNEMIEKE VERBEKEN*
"7 rue des Coquilles, 67500 Haguenau, France
? Department of Biology, Ghent University,
K. L. Ledeganckstraat 35, 9000 Ghent, Belgium
CORRESPONDENCE TO: *mieke. verbeken@ugent.be 'jmtrendel@free.fr *felix.hampe@email.de
ABSTRACT—Based on morphological, molecular, and ecological data, a new species of
Russula sect. Russula, found on several occasions under deciduous trees in Northern Alsace
is described and illustrated as: Russula vinosoflavescens, belonging to R. subsect. Sardoninae.
Key worps—Basidiomycota, Russulaceae, ITS, Russula persicina, phylogeny
Introduction
Russula Pers. (Russulales, Basidiomycota) is an ectomycorrhizal genus with
a world-wide distribution that is particularly well studied in temperate regions,
notably in Europe (Romagnesi 1967; Sarnari 1998, 2005). The ectomycorrhizal
associations, which often involve a selective fungus-plant partnership, were
sometimes advantageously used to delimit infrageneric taxa. Such has been
the case for R. subsect. Sardoninae Singer (Sarnari 1998, emend.), which is
divided into series—provisionally defined by Sarnari (1998)—comprising, on
one hand, species strictly associated with conifers (R. ser. Sardonia and R. ser.
Sanguinea, both essentially corresponding to R. subsect. Sardoninae as
conceived by Romagnesi 1967, 1987) and, on the other hand, species associated
with deciduous trees, sometimes in a host-specific relationship (R. ser.
Exalbicans = R. subsect. Exalbicantinae Singer as retained by Romagnesi, with
species found only under birches) and sometimes with a broader host-range
(R. ser. Persicina = R. subsect. Persicinae Romagn., species that can be found
under oaks, hornbeams, beeches, birches, chestnuts, poplars, willows).
708 ... Trendel, Hampe & Verbeken
TABLE 1. Russula, Gymnomyces, and uncultured specimens with GenBank
and UNITE sequence accession numbers used in the molecular analyses.
SPECIES
G. gilkeyae
G. monosporus
root sample (uncultured)
R. americana
R. aquosa
R. atrorubens
R. bresadolae
(“R. atropurpurea”)
R. cavipes
Russula cf. “ammodica”
R. citrinochlora
R. consobrina
R. depallens
R. emetica
R. exalbicans
R. fageticola (“R. mairei”)
R. fageticola (“R. nobilis”)
R. fellea
R. fragilis
R. gracillima
R. luteotacta
R. ochroleuca
R. persicina
R. pumila
R. queletii
SPECIMEN
JT 2572
OSC 117360
F 18871
TU 101831
TU 101708
TU 101691
TU 101851
HUE 178
PRM 858109
RT 9149
HUE 163
F 18874
TU 101905
20024
TU 118108
TU 106965
TU 101829
TU 101838
F 14309
DG 18
DG 44
2-11171S76
FH 12262
HUE 054
HUE 218
HUE 177
UE 2006-11-08-22
UE 2004-20-09-04
TU 101725
TU 106437
TU 106379
FH 12-187
TU 118113
LW 115
TU 101826
TU 106950
F 133
L 3X87
FH 12-237
RT 5140
COUNTRY
USA
USA
Estonia
Estonia
Estonia
Estonia
Germany
Czech Republic
Italy
Germany
USA
Norway
Sweden
Estonia
Estonia
Estonia
Estonia
United Kingdom
United Kingdom
Germany
Germany
Germany
Germany
Italy
Sweden
Estonia
Estonia
Estonia
Germany
Estonia
Germany
Estonia
Estonia
Finland
Germany
Italy
SEQUENCE
AY239346
EU669222
JQ890299
JN197654
EF434062
GU997950
GU997948
HQ604839
UDB015988
UDB011290
UDBO011358
UDB016006
UDB000313
HG423574
JF908681
AF418623
HQ604840
UDB016045
UDB002550
UDBO011223
UDBO011207
UDB015986
UDB015994
AY228360
JQ888196
UDB001628
AY061674
KT934013
UDB000346
UDB000345
UDB000314
UDB018436
UDB018434
UDBO011361
UDBO011176
UDBO011166
KT933991
UDBO011228
UDB000295
UDB015984
UDBO011196
UDB009582
FM993279
KT934007
JF908668
Russula vinosoflavescens sp. nov. (France) ... 709
R. renidens TU 101809 Estonia UDB015975
JR1758F™ Finland UDBO011117
R. rhodomelanea FH 2011-BT307 Germany UDB018429
IB 92/451 #7 Italy UDB018435
R. sanguinea TU 106710 Estonia UDB019728
FH 12240 Germany KT934008
R. sardonia TU 106951 Estonia UDBO011197
FH 12215 Germany KT933999
R. silvestris JK 11080504 Sweden UDBO018431
JK 11080802 Sweden UDB018430
R. thindii™™ BSHC KD11-095 India KM386693
R. torulosa TU 106444 Estonia UDBO011177
TU 101626 Italy UDB016261
R. vinosoflavescens JMT 05081006 France UDB031189
JMT 08081512 France UDB031188
JMT 14090502 # France UDB031187
JMT 05081007 France UDB031190
JMT 10080701 France UDB024104
JMT 13090715 France UDB024105
Outgroup
R. puellula (“R. puellaris”) MC 01-502 Denmark UDB000010
R. cuprea FH 12250 Germany KT934010
R. olivobrunnea TU 101883 Finland UDB016034
HT holotype; ™ isotype.
The aim of this paper is to introduce a new Russula species occurring in
deciduous forests of Northern Alsace (NE France) and belonging to R. subsect.
Sardoninae. Our initial morphological concept of the species, based on several
collections from various localities, is fully supported by molecular data. ITS
sequence analysis shows that the different collections form a well-supported
clade in the Russula phylogenetic tree.
Materials & methods
Sampling
All collections of the new taxon were made by Jean Michel Trendel and are deposited
in GENT Herbarium. These collections are referred to with an eight-digit code
YYMMDDxXxX (year; month; day; collection number). All collecting sites are located
in Northern Alsace (NE France); the code of the municipality is that assigned by the
National Institute for statistics and economic studies (INSEE); names (in quotation
marks) of forest or place are those indicated on the topographic maps (1:25000 scale)
from the Institut national de linformation géographique et forestiére (IGN); map
coordinates of the collections refer to the Universal Transverse Mercator (UTM)
32N kilometer grid system; geological data (underlying rock) were mainly obtained
710 ... Trendel, Hampe & Verbeken
from geological maps (1:50000) edited by the Bureau de Recherches Géologiques et
Miniéres (BRGM).
Morphological analysis
Specimens were photographed in situ. The pictures are available at
http://www2.muse.it/russulales-news/. Microscopic observations were carried out
on fresh material. Pileipellis elements were studied on radial sections made by hand
midway from the cap margin, mounted either in SDS Congo red (Clémengon 1999)
or in distilled water. The search for acido-resistant encrustations was performed by
staining with Ziehl carbol fuchsin followed by a rapid differentiation using a 1M HCl
solution. Pileocystidia content was revealed using sulfovanillin (SV) freshly prepared
with a 50% or 80% H,SO, aqueous solution. Hymenial elements were studied in the
above-mentioned observation media. In some cases, additional controls were carried
out on very fine cap sections made from dried material and further rehydrated in a
moisture-saturated closed chamber for 24 h before observation in water. Basidiospores
(from spore deposits) were examined in Melzer’s reagent and spore measurements
(ornamentation excluded) were recorded randomly for 300 spores (6 collections) in
side view. Measurements are given as (MINa) [AVa - 2*SD]—Ava-AVb-[AVb + 2*SD]
(MAXb), with AVa = lowest mean value for the measured collections and MINa the
minimum value corresponding with this mean value, AVb = greatest mean value and
MAXb the maximum value corresponding with this mean value, and SD = standard
deviation calculated for the measurements of one collection (minimum and maximum
value are only given if not in the 2*SD-interval). Q stands for spore “length/width ratio”
and is given as (MINQa)-Qa-Qb-(MAXQb) with Qa and Qb being the lowest, and the
highest respectively, mean ratio for the measured specimen. The colour of the spore
deposit, referring to Romagnesi’s (1967) scale, was assessed against a personal chart
(JMT) and the Dagron’s chart (unpublished). Macrochemical reactions were determined
using FeSO, in crystalline form, a 4% phenol solution, and a strong Guaiac solution
made of permanently soaked Guaiac wood in 95% ethanol.
Molecular analysis
Total genomic DNA was extracted from dried material according to Nuytinck &
Verbeken (2003), with modifications described in Van de Putte et al. (2010). The ITS
region was amplified using the primers ITS1F-ITS4 (White et al. 1990, Gardes & Bruns
1993) and with polymerase PerfectTAQ (5 PRIME, Hilden, Germany) in accordance
with the manufacturer's recommendation. PCR amplification followed Eberhardt
(2012), and the PCR products were purified using the Qiaquick PCR Purification
Kit (Qiagen, Hilden, Germany) and directly sequenced with BigDye 3.1 technology
(Applied Biosystems, now Thermo Fisher Scientific, Wilmington, USA). Specimens
JMT-05081006, JMT-05081007, and JMT-05081512 were extracted and sequenced in
Muséum National d'Histoire Naturelle de Paris (Marc-André Selosse) following Séne
et-al7(2015).
Raw sequences were edited in the BioEdit Sequence Alignment Editor version 7.2.5
(Hall, 2013) or Sequencher version 4.8 (Gene Codes Corporation). Edited sequences
Russula vinosoflavescens sp. nov. (France) ... 711
were aligned by MAFFT version 7 using the strategy E-INS-i (Katoh & Standley 2013).
Maximum-likelihood searches for tree building were carried out locally with 100
replicates with the GTR+GAMMA model, selecting the best solution of all replicates
analysis in RaXML 8.1.12 (Stamatakis 2014). Fast bootstrap searches were done locally
or through the CIPRES Science Gateway (Miller et al. 2010) with 10 000 replicates.
The final alignment included a total number of 67 ITS sequences, with 21 sequences
corresponding to the current concept of R. subsect. Sardoninae (Sarnari 1998). Except
for the R. vinosoflavescens collections cited as studied material all sequences were
retrieved from GenBank or UNITE (TABLE 1). The tree with the highest log likelihood
(-—4523.1303) is shown. Initial tree(s) for the heuristic search were obtained automatically
by applying the Maximum Parsimony method.
Taxonomy
Russula vinosoflavescens Trendel & F. Hampe, sp. nov. Figures 1, 2
MycoBAnk MB 819428
Differs from Russula persicina by its colour range, its mild or only slightly acrid taste,
its reticulate spore ornamentation, and the presence of granular pigments in cuticular
hyphae.
Type: France, Alsace, Morsbronn-les-Bains (67303), “Niederwald’, UTM 32N:
0406318/5419158, 213 m alt., on argillaceous soil with eumull humus (underlying
rock: upper Keuper red marls), under Quercus robur and Carpinus betulus with some
Fagus sylvatica nearby, in an area characterized by calcicolous mycoflora with numerous
Phlegmacium (Cortinarius) species, 5 Sept. 2014, JMT-14090502 (Holotype, GENT;
UNITE UDBo31187).
EryMoLocy: Referring to the purplish-red wine-coloured (vinoso) cap and the
tendency of the cap to become brown-yellowish or ochraceous (flavescens) as well as
when handling the stipe.
PitEus 4.0-6.5 cm diam., more or less fleshy, rather firm to very firm but with
margin occasionally slightly elastic, often irregular, at times knotty, convex-
flattened with the margin sometimes incurved or even inrolled, then broadly
but shallowly depressed; margin not or shortly sulcate, often flexuose-undulate
or lobed, in some cases showing a very clear, whitish border extending to the
edge of the gills (giving a “festooned” appearance); surface rugulose, more or
less radially veined, frequently uneven-bumpy in the centre, a little shiny in wet
conditions, but soon dry (except sometimes in the centre which remains a little
greasy) and then more mat, purple-red, carmine-red, vinaceous-red, somewhat
violet, rather vivid, but frequently paler, livid pinkish-vinaceous, with mauve
(lilac) greyish shades that can entirely replace the reddish tinges, even more
clearly greyish purple at the margin, becoming yellowish cream-ochraceous
from the centre or showing locally ochre-bistre areas merging in the reddish
and greyish mauve tints, sometimes becoming almost totally beige-brown,
712 ... Trendel, Hampe & Verbeken
ochreous-brown, with livid purple-violet glints mostly near the margin and a
residual purple-brown at centre. LAMELLAE adnate or attenuated-subdecurrent,
sometimes sub-emarginate, with a variable number of lamellulae, crowded to
rather spaced, interveined, anastomosing and more or less forked at every
level, in some cases more particularly near the stem (exceptionally regular,
without any forking or anastomosing), (sub)acute at the cap margin (rarely
subobtuse), narrow (0.3-0.5 cm, exceptionally broader <0.7 cm), whitish or
very pale cream; edge entire but often somewhat irregularly wavy. STIPE 2-6 x
0.9-1.5(-2) cm, cylindrical or slightly enlarged downwards, usually rounded
at the base (in a few cases tapering), sometimes eccentric, firm to almost hard,
with a firm medulla (exceptionally hollow-fistulous); surface nearly smooth,
finely wrinkled, white with occasionally a purple tinge (in one case remarkably
grey-purplish, somewhat brownish), distinctly yellowing (dirty, often with
greyish-brown shades) when handled or bruised. CONTEXT white, in the pileus
almost unchanging, in medulla of stipe slightly yellowing, sometimes a little
purple-violet beneath the cuticle in the most coloured forms; taste mild (or
almost mild, occasionally with an unpleasant aftertaste, somewhat bitter),
mild to slightly (and sometimes volatile) acrid in the gills, exceptionally more
strongly acrid; smell not distinctive.
MACROCHEMICAL REACTIONS FeSO: rose-orange; Phenol: brown; Guaiac:
in most cases reacting on the stem surface almost immediately (<2 s, in one
case more slowly), but of little intensity at the beginning, developing (10-15 s)
in blue of medium intensity, exceptionally with a stronger reaction, (dark blue);
on gills, the reaction developing slowly and weakly (blue-green to light blue).
SPORE PRINT cream, II(a)b-c.
Basip1osporss subglobose to broadly ellipsoid, 6.4-7.2-8.1-9.0 x 5.9-6.5-
6.9-7.7 um (n = 300), Q = 1.06-1.11-1.20-1.30; ornamentation amyloid,
often with incompletely amyloid warts; warts conical-obtuse, or more acute,
sometimes somewhat truncated, generally around 0.4-0.8 um high, but
also frequently reaching 1.0 um or higher, sometimes locally catenulate,
interconnected by crests or more frequently by (often very fine) connectives,
forming a well-developed and often nearly complete (with numerous meshes)
reticulum, which may appear confused in some cases; suprahilar plage
moderately amyloid, more or less bordered by small warts or low crests.
Basip1A 4-spored, 42-57 x 9.5-12.5 um, including sterigmata (about 5 um
long). PLEUROMACROCYSTIDIA 60-90 x 10.5-14.5 um, numerous, fusiform
or more or less clavate, most commonly with a pointed appendage, which
can show constrictions, more rarely ending in a short obtuse protuberance,
Russula vinosoflavescens sp. nov. (France) ... 713
i a, Mga
Fic. 1. Russula vinosoflavescens: colour forms observed. A, B. holotype, collection JMT-14090502;
C. collection JMT-05082042, with marked reddish pigmentation; D. collection JMT-05081006,
with marked reddish pigmentation; E. collection JMT-08081512, with predominating ochraceous
colour; EF. collection JMT-10080701, with mauve-lilac colour reminiscent of a Griseinae.
714... Trendel, Hampe & Verbeken
strongly staining with SV. PILEIPELLIS composed of hyphae containing reddish-
violaceous granular pigments; epicuticular elements (2.0-—)2.5-4.0 um diam.,
sometimes branched, flexuose, sometimes inflated up to 5.5 um; terminal cell
obtuse, or slightly thickened with a short appendage, also subcapitulate, or
(exceptionally) pear-shaped <7.0 um diam., but also conversely more or less
attenuated (sometimes abruptly). PILkeocystTip1a of two types; type 1 usually
unicellular (occasionally with 1 septum), rarely bifid, clavate—-fusiform, less
frequently cylindric, 4.0-9.5 um diam., often appendiculate-capitulate or
showing an obtuse apical protuberance, sometimes slender, (at least 120 um
long, narrowed at the base to 2.5-3.0 um diam.), with a content reacting variably
depending on the collection—but also on a cuticular section—when treated
with SV, staining (rather) strongly (grey-blackish) or weakly (grey-pinkish)
or even remaining inert (pinkish); type 2 lactiferoid (some clearly termini of
ascending lactifers), pluriseptate, more or less regularly cylindrical, 6.0-10.5
uum diam., with variably shaped terminal elements that sometimes taper (with
possible constrictions), sometimes are somewhat thickened or subclavate, or
with type 1 type appendages, originating in the deeper pellis, distinctly reacting
with SV (except in one specimen where the granular content of pileocystidia
type 2 is inert or reacts only very weakly in SV); without encrusted elements
after treatment with Ziehl fuchsin.
ECOLOGY & DISTRIBUTION—Under deciduous trees, associated with
Carpinus betulus, or Quercus (Q. robur or Q. petraea), or both (possibly also
with Fagus sylvatica), in a rather wet environment (possibly periodically drier),
on more or less argillaceous soils, neutral to superficially slightly acidic, usually
nutrient-rich with a rather high base saturation level (calcium-rich), with
mesotrophic to eutrophic mull humus. Phenology: late July to early September
(summer fruiting). Known only from six sites (9 collections) in Northern
Alsace (France).
ADDITIONAL SPECIMENS EXAMINED—FRANCE, ALSACE. Bas-Rhin, Mattstall,
commune associated with Lembach (67263), ‘Sauerhald} UTM 32N: 0409918/5426543,
alt. 203 m, underlying rock: Upper Muschelkalk marly calcareous formation, under
Quercus, Carpinus, Fagus (forest locally rich in orchids, e.g., Epipactis microphylla),
JMT-13090715 (GENT, UNITE: UDB024105); Dauendorf (67087), “Herrenwald}
0402753/5410066, 168 m, Loess from the Quaternary (and more ancient alluvial
deposits?), in a wet environment, nitrogen-rich and with a high base saturation level,
under Carpinus, Quercus, Alnus glutinosa, Fraxinus excelsior, Ulmus sp., and with a
herbaceous layer comprising Allium ursinum, Carex sylvatica, Circaea lutetiana, Geum
urbanum, Glechoma hederacea, Paris quadrifolia, Stachys sylvatica, JMT-10080701
(GENT, UDB024104); 0402762/5409965, 171 m, Loess from the Quaternary (and
more ancient alluvial deposits?), under Carpinus, Quercus, JMT-05082042 (GENT);
Russula vinosoflavescens sp. nov. (France) ... 715
Fic. 2. Russula vinosoflavescens (holotype, collection JMT- 14090502). A. fruitbody in sectional
view; B. pileus epicutis: pileocystidia (with content shown schematically) and hyphal terminal
elements; C. hymenial cystidia on gill sides; D. spores as observed in Melzer’s reagent. Scale bars:
A=1cm;B=10 um; C = 20 um; D=5 um.
716 ... Trendel, Hampe & Verbeken
Bettwiller (67036), “Buchwald, 0365394/5416535, 347 m, Upper Muschelkalk marly
calcareous formation, under Quercus, Carpinus, with some Fagus and Prunus avium,
JMT-05081006 (GENT, UDB031189) and JMT-05081007 (GENT, UDB031190), most
likely belonging to the same mycelium; Forstheim (67141), ‘der Wald; 0405631/5415509,
211 m, Pliocene sands and clays, wet environment, under Quercus, Carpinus, Populus
tremula with Russula lutensis Romagn. and R. rutila Romagn., JMT-08081512 (GENT,
UDB031188); Drusenheim (67106), ‘Barrwald, approximately 0420900/5401000, 124 m,
Quaternary alluvial deposits, wet environment, on bare blackish ground, with Carpinus,
Quercus, Alnus glutinosa, Fraxinus excelsior, 25/07/1981, JMT-81072501 (GENT, spore
print only available).
Discussion
Molecular analysis
The six ITS sequences of the newly described R. vinosoflavescens form
an independent clade, which receives high bootstrap support (Fic. 3). The
clade corresponding to R. vinosoflavescens is nested within a large, supported
(bootstrap value 75) clade that includes the typical European representatives of
R. subsect. Sardoninae (R. sardonia Fr., R. queletii Fr., R. torulosa Bres.). Beside
these, it contains a number of species (e.g., R. cavipes Britzelm., R. exalbicans
(Pers.) Melzer & Zvara, R. gracillima Jul. Schaff., R. luteotacta Rea, R. persicina
Krombh., R. renidens Ruots. et al., R. sanguinea Fr.) included by Sarnari (1998)
in his emended concept of R. subsect. Sardoninae but attributed to different
(although closely related) infrageneric taxa by previous authors (Romagnesi
1967, Singer 1986, Bon 1988). In addition to the abovementioned species,
which fairly well represent R. subsect. Sardoninae sensu Sarnari, the clade
includes extra-European Sardoninae species such as R. thindii K. Das & S.L.
Miller and some poorly known species with an uncertain systematic position
(e.g., R. americana Singer, R. citrinochlora Singer).
The deeper phylogenetic relationships within the Sardoninae clade as
well as its external relationships with R. fellea (Fr.) Fr., R. consobrina (Fr.) Fr.,
R. ochroleuca Fr., and the large clade representing R. subsect. Russula
s.l. (including a sequence of the generic type R. emetica (Schaeff.) Pers.)
remain poorly resolved in the ITS analysis. Within the Sardoninae clade,
R. vinosoflavescens occupies a rather isolated position in a weakly supported
subclade containing R. renidens and samples identified as R. citrinochlora.
Morphological analysis
Despite its mild taste (in most cases only weakly acrid in gills),
R. vinosoflavescens shows a combination of characters—a more or less fleshy
firm pileus, cuticle often rugulose with polychromic colours, cream spore
Russula vinosoflavescens sp. nov. (France) ... 717
99 , KT934007 R. queletii
98 JF908668 R. queletii
UDB011177R. torulosa
100 ' UDB016261 R. torulosa
KM386693 R. thindii HOLOTYPE
HQ604839 R. americana
100 ' HQ604840 R. cf. ammodica
17 UDB019728 R. sanguinea
100 'KT934008 R. sanguinea
UDB011197 R.sardonia
100 & KT933999 R. sardonia
JQ890299 root sample
JF908681 R. cavipes
75 'AF418623R. cavipes
100 ; UDB011166R. luteotacta
KT933991 R. luteotacta
100 , UDB015984 R. persicina
UDB011196 R. persicina
100 ; UDB011361 R. gracillima
98 UDB011176R. gracillima
UDB015986 R. depallens
a8 AY061674R. exalbicans
9° 1 UDB015994 R. depallens
100 ; EU669222 G. monosporus
AY239346G. gilkeyae
98 , UDB016045 R. citrinochlora
UDB002550R. citrinochlora
UDB015975R. renidens
JN197654 root sample
UDB011117R.renidens ISOTYPE
EF434062 uncultured fungus ALASKA
100 GU997950 rootsample
GU997948 rootsample
80 UDB031188 - JMT-08081512
UDB031189 -JMT-05081006
UDB031187-JMT-14090502HOLOTYPE \ Russula
99 | UDB031190 - JMT-05081007 vinosoflavescens
UDB024104 - JMT-10080701
UDB024105 - JMT-13090715
UDB000345R. fellea
100 'UDB000314 R. fellea
100 , UDB011223 R. consobrina
UDB011207 R. consobrina
100 - UDB015988 R. aquosa
90 UDB011290 R. aquosa
100 ; KT934013 R. fageticola
93 UDB000346 R. fageticola
90 ; UDB011358 R. atrorubens
17 98 UDBO016006R. atrorubens
AY 228360 R. emetica
97 | UDB001628 R.emetica
JQ888196 R.emetica
100 } UDB018431 R. silvestris
UDB018430 R. silvestris
98 - UDB018436R. fragilis
UDB018434 R. fragilis
62 UDB018429 R. rhodomelanea
100 !' UDB018435 R. rhodomelanea HOLOTYPE
100 } UDB011228 R. ochroleuca
UDB000295 R. ochroleuca
99 ; UDB009582 R. pumila
FM993279 R. pumila
65
94
UDB000313R. bresadolae
100 'HG423574 R. bresadolae
UDB000010R. puellula
100 KT934010R. cuprea Outgroup
0.05 98 UDB016034 R. olivobrunnea
Fic. 3. Molecular phylogenetic analysis by Maximum Likelihood method-based tree with the
highest log likelihood (—4523.1303), based on ITS sequences.
718 ... Trendel, Hampe & Verbeken
print, well-characterised and non-encrusted pileocystidia—that is consistent
with its position in R. subsect. Sardoninae sensu Sarnari. Moreover, by its
habitus, somewhat decurrent narrow gills, and clear yellowing of its stipe when
bruised, this species is very reminiscent of certain forms of R. persicina. Russula
vinosoflavescens differs, however, unambiguously from the latter by its colours,
which are never ‘pure’ red. Indeed, even in its reddest forms, the cap always
displays some purple-vinaceous tinges, presumably reflecting the existence of
one or several blue pigments, occurring in sufficient amount to significantly
shade the red ones. The presence of these blue pigments becomes even more
apparent when grey-mauve (lilac) tints replace—sometimes completely—the
reddish colouring. In addition, this russula appears to fade readily to yellowish
ochre, or even brown-beige, and therefore colour forms devoid of any reddish
or bluish shades might be expected.
Other striking features are the often rugulose surface of the cuticle and the
strong tendency of the lamellae to become anastomosed-forked, though it
should be kept in mind that these characters are variable. The same is true for
the variability of the speed and intensity of the Guaiac reaction, which is mostly
in an average position on the reactivity scale.
One of the most striking microscopical features is the aggregation of
pigments into granules (sometimes distinctly combining red and blue
pigments) that are generally easily observed in the cap cuticular hyphae. The
cuticular structure of specimens for which we were unable to examine in fresh
condition was re-examined in dried material. Concerning the least pigmented
collection JMT-08081512 (Fic. 1-E), cuticular sections from the central part
of the cap—apparently the richest in residual red and blue pigments—revealed
the presence of pigmented grains. Our search was much less conclusive for the
vividly coloured specimens JMT-05081006 (Fic. 1-D), and even if we were able
to show the presence of sparse coloured granules, it appears that a significant
amount of pigment is either vacuolar (soluble) or at least not distinctly granular.
In addition, examination of the cuticle of collection JMT-05082042 (Fic. 1-C),
not studied in a fresh state, clearly revealed the presence of pigmented granules.
As reported by Romagnesi (1967) and Sarnari (1998), granular pigments are
highly characteristic of R. subg. Heterophyllidia but are also found occasionally
in some species belonging to other groups. In this respect, one should especially
mention R. renidens, a Nordic russula growing under birches, which shows
abundant granular red-purplish pigments, the latter probably co-occurring
in a soluble form accounting for the intensity of its colours (Sarnari 1998).
A similar situation may also exist for some of our vividly coloured collections
Russula vinosoflavescens sp. nov. (France) ... 719
(notably JMT-05081006 and JMT-05081007). Indeed, it seems likely that
pigment aggregates contribute less efficiently to global colouring compared to
those in solution, and that a strong predominance of granular pigments over
their soluble counterparts may explain the relative paleness of some of our
collections.
The morphology of the terminal hyphal cells of the epicuticular elements
appears too variable to be usable for species recognition. On the other hand,
pileocystidial shape seems informative at species level. It is also significant that
this characteristic can be found in almost all members of R. subsect. Sardoninae
as defined by Sarnari, where the new species belongs.
Another important diagnostic feature is the strongly reticulate spore
ornamentation that distinguishes R. vinosoflavescens from similar species
sharing the same cream spore print and also growing under deciduous trees,
such as R. persicina or the birch associates, R. exalbicans and R. renidens.
Ecology
Within Russula subsect. Sardoninae, partitioned by Sarnari (1998) into
ecological series based on Russula/tree partnerships, R. vinosoflavescens fits in
R. ser. Persicina, devoted to species associated with broadleaf trees (but not
strictly with birches); the other series cover species linked strictly to either
conifers (R. ser. Sardonia and R. ser. Sanguinea) or birches (R. ser. Exalbicans).
Indeed, R. vinosoflavescens appears associated with oak (Quercus robur or
Q. petraea) or hornbeam (Carpinus betulus) or both, the only trees present
on all collecting sites. A wider host association might include beech (Fagus
sylvatica, generally well represented in the forests of interest but sometimes
rather far away from the fruiting bodies) or, less likely, aspen (Populus tremula,
which can occur in more complex environments). Birch (Betula spp.), absent
from all the prospected sites, should not be considered a potential associate.
However, regarding Romagnesi’s (1967) initial concept of Persicinae, which
refers to “intermediate forms between Emeticinae and Sardoninae [both sensu
Romagnesi!], with less reticulate spores and gills more decurrent or weeping
than the former, and more purely red in colour than the latter” [a concept at
least partly shared by Sarnari (1998)—“pileus pure red”], it becomes necessary
to restrict Sarnari’s R. ser. Persicina definition exclusively to ecology, as in
R. vinosoflavescens colouration is not restricted to pure red and its spores are
rather strongly reticulate. In his classification, Bon (1988) does not retain
Romagnesi’s Persicinae group but treats the different taxa of R. persicina in his
emended R. subsect. Exalbicantinae. In this context, R. vinosoflavescens would
take its place beside R. exalbicans, which also often displays washed-out greyish
720 ... Trendel, Hampe & Verbeken
colours, and R. renidens, genetically a very closely related species sharing some
outstanding morphological features such as pigments that are at least partly
granular or lamellae that are more or less forked.
Acknowledgments
We are extremely thankful to Josie Lambourdiere and Prof. Marc-André Selosse
(Muséum national d’Histoire Naturelle, Paris) for some additional molecular analyses,
and to Dr. Philippe Schaeffer for helpful discussions. Dr. Ursula Eberhardt and Geoffrey
Kibby are acknowledged for useful comments and reviewing the paper.
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Clémencon H. 1999. Vom Umgang mit Kongorot. Schweiz. Z. Pilzk. 77(5): 247-250.
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(eds). DNA Barcodes: Methods and Protocols, Methods in Molecular Biology, vol. 858. New
York, Humana Press. https://doi-org/10.1007/978-1-61779-591-6_9
Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes -
application to the identification of mycorrhizae and rusts. Mol. Ecol. 2(2): 113-118.
https://doi.org/10.1111/j.1365-294X.1993.tb00005.x
Hall T. 2013. BioEdit, Biological sequence alignment editor for Windows 5/98/NT/2000/XP/7,
version 7.2.5. Ibis Biosciences, Carlsbad. https://www.mbio.ncsu.edu/bioedit/bioedit.html
(accessed 15 March 2016).
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improvements in performance and usability. Mol. Biol. Evol. 30(4): 772-780.
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(GCE), 14 Nov. 2010, New Orleans, Louisiana. https://doi.org/10.1109/GCE.2010.5676129
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morphological analyses. Mycol. Progress 2(3): 227-234.
https://doi.org/10.1007/s11557-006-0060-5
Romagnesi H. 1967. Les Russules d’Europe et d'Afrique du Nord. Paris, Bordas.
Romagnesi H. 1987. Statuts et noms nouveaux pour les taxa infragénériques dans le genre Russula.
Doc. Mycol. 18(69): 39-40.
Sarnari M. 1998. Monografia illustrata del Genere Russula in Europa, vol 1. Vicenza, Associazione
mycologica Bresadola (AMB), Fondazione Centro studi micologici.
Sarnari M. 2005. Monografia illustrata del Genere Russula in Europa, vol 2. Vicenza, Associazione
mycologica Bresadola (AMB), Fondazione Centro studi micologici.
Séne S, Avril R, Chaintreuil C, Geoffroy A, Ndiaye C, Diédhiou AG, Sadio O, Courtecuisse R, Sylla
SN, Selosse MA, Ba A. 2015. Ectomycorrhizal fungal communities of Coccoloba uvifera (L.) L.
mature trees and seedlings in the neotropical coastal forests of Guadeloupe (Lesser Antilles).
Mycorrhiza 25: 547-559. https://doi.org/10.1007/s00572-015-0633-8
Singer R. 1986. The Agaricales in modern taxonomy. Konigstein, Koeltz Scientific Books.
Stamatakis A. 2014. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large
phylogenies. Bioinformatics 30(9): 1312-313. https://doi.org/10.1093/bioinformatics/btu033
Russula vinosoflavescens sp. nov. (France) ... 721
Van de Putte K, Nuytinck J, Stubbe D, Le HT, Verbeken A. 2010. Lactarius volemus sensu lato
(Russulales) from northern Thailand: morphological and phylogenetic species concepts
explored. Fungal Diversity 45(1): 99-130. https://doi.org/10.1007/s13225-010-0070-0
White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal
ribosomal RNA genes for phylogenetics. 315-322, in: MA Innis et al. (eds). PCR Protocols:
a guide to methods and applications. San Diego, Academic Press.
https://doi.org/10.1016/b978-0-12-372180-8.50042-1
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 723-744
https://doi.org/10.5248/132.723
Triadelphia acericola and T. centroseptata spp. nov.,
and a synopsis of the genus
DeE-WEI LI »** & JIAN-REN YE ”?
"The Connecticut Agricultural Experiment Station Valley Laboratory,
153 Cook Hill Road, Windsor, CT 06095, USA
?Co-Innovation Center for Sustainable Forestry in Southern China & ° College of Forestry,
Nanjing Forestry University, Nanjing, Jiangsu 210037, China
* CORRESPONDENCE TO: dewei.li@ct.gov
ABSTRACT—A new hyphomycete species found during a collection of microfungi from
plant debris and decaying wood in Connecticut, USA, is described and illustrated here
as Triadelphia acericola. The new species produces four conidial forms: form (a), the
predominant form, is cylindrical and 2-septate, with the apical septum covered by a wide
dark band; form (b) is oblong or ellipsoidal and 1-septate; form (d) is clavate and pluriseptate,
with the middle transverse septum covered by a dark band; and form (f) is globose. Literature
on all published Triadelphia names is reviewed, and a comparative synopsis of the genus and
a key to Triadelphia species are provided. “Triadelphia centroseptata” nom. inval. is validated
by designation of a holotype, and T: archontophoenicicola nom. nov. is proposed to replace the
illegitimate homonym, T: australiensis Joanne E. Taylor et al.
Key worps—Acer, anamorph, nomenclature, pleomorphic, saprobe
Introduction
Triadelphia was erected by Shearer & Crane (1971) as a monotypic genus
typified by T. heterospora. The generic concept of Triadelphia was originally
described as: “Conidiogenous cells subspherical, subhyaline to dark colored,
producing conidia from a very limited area at the apex. Conidia readily
deciduous, leaving a pore when detached, cylindrical, rounded at the apex,
rounded to slightly truncate at base, dark colored, and septate.” (Shearer &
Crane 1971). Triadelphia heterospora develops two different forms of conidia: 1)
cylindrical and 2-septate and 2) broadly obclavate to ellipsoid and 4—7-septate;
724 ... Li & Ye
both conidial forms have at least one septum covered by a dark band (Shearer
& Crane 1971).
Hughes & Pirozynski (1973) transferred Dicoccum inquinans to Triadelphia.
Maggi et al. (1978) described two new species: T: loudetiae and T. pulvinata.
Constantinescu & Samson (1982), who studied herbarium specimens of
T. inquinans, type specimens of T: heterospora, and ex-type cultures of
T: loudetiae and T. pulvinata, found those species to be pleomorphic, developing
2-5 forms of conidia, and emended the generic description of Triadelphia to:
“Conidiogenous cells arising from undifferentiated hyphae, hyaline or pale
brown, flask-shaped, fusiform, cylindrical or clavate, solitary or agglomerated in
sporodochium-like structures, mostly with determinate growth and producing
one apical conidium. Conidia blastic, of at least 2 (sometimes 5) forms in every
species: (a) cylindrical, brownish, 1-2 septate; (b) clavate, dark brown, unispetate
[sic]; (c & d) obclavate to acicular with a narrow long tip, hyaline or yellowish
brown, multiseptate; (e) allantoid, hyaline or pale yellowish, 0-3-septate,
sometimes also (f) obovate to broadly ellipsoidal, pale brown, unicellular.”
Constantinescu & Samson (1982) simultaneously described a new species,
T. romanica, and transferred Stemphyliomma alabamensis to Triadelphia.
Kirk (1983) transferred Sporidesmium uniseptatum to Triadelphia based on
conidiogenous cell morphology and conidial ontogeny and morphology, which
were all similar to the previously published six Triadelphia species. Mercado
Sierra & Castafieda-Ruiz (1983) published T! stilboidea, a synnematous species
from Cuba, and Révay (1987) published T: hungarica from Hungary. Tzean &
Chen (1989) described as new T. diversa; they also reviewed and compared
eight Triadelphia species but did not include T. stilboidea or T: hungarica in
their review and key.
Sutton (1989) described Triadelphia australiensis, a species with only one
conidial form, although he acknowledged that all other Triadelphia species
develop at least two forms of conidia. Matsushima (1989, 1995) described two
new species, T: queenslandica and T. synnematofera (the second synnematous
species in Triadelphia). Révay (1993) published her second new species,
T. morgoensis, with a key to 12 species; she also noted that as T. stilboidea
probably did not belong in Triadelphia due to its synnematous nature, she
omitted Triadelphia stilboidea from her key.
Venkateshwarlu et al. (1997) added T: centroseptata to the genus.
Manoharachary et al. (2001) described T. corticola as having two conidial
“forms, which appear to represent the same form varying only in conidial
width and shape: conidia in T’ corticola share the same length, septation
Triadelphia acericola & T. centroseptata spp. nov. ... 725
characters, and conidiogenesis and belong to form (a) of Constantinescu &
Samson (1982). The most recent species added to Triadelphia is T: australiensis
Joanne E. Taylor et al. (Taylor & Hyde 2003). Of the eighteen species described
in Triadelphia, “T. centroseptata” was invalidly published and is validated here.
Since the morphological characters and conidium ontogeny of our specimen
collected in Windsor, Connecticut, cannot be ascribed to any previously
described Triadelphia species, we propose it here as a new species, T. acericola.
Materials & methods
An undescribed hyphomycete was collected from a piece of inner bark of Acer
palmatum during a collection of fungi from plant debris and decaying wood in a mixed
forest, wood log piles, and a landscape garden at the Valley Laboratory, Windsor, CT, in
June 2013. The specimen was placed in a paper bag and brought back to the laboratory.
Isolation on to MEA and CMA was attempted in the same day. However, isolation was
not successful. Therefore a small scalpel was used to remove fungal structures directly
from the samples for morphological observation under an Olympus BX40 compound
microscope. Conidiophores, conidia, and other fungal structures were mounted in
85% lactic acid or lacto-fuchsin (0.1 g acid fuchsin, 100 ml 85% lactic acid; Carmichael
1955). The fungal structures were microscopically observed and measured under
Nomarski differential interference contrast optics. Photomicrographs were captured
using an Olympus Microfire digital camera (Goleta, CA). Data of the fungal structure
measurements were statistically analyzed using Microsoft Office Excel 2013 with 95%
confidence interval of means. The results were presented as ranges and mean + standard
deviation as well as Q (ratio of conidial length/width).
Taxonomy
Triadelphia acericola D.W. Li, sp. nov. FIGS 1, 2A
MycoBank MB813008
Differs from Triadelphia heterospora, T. inquinans, and T. diversa by not having conidial
forms (c) and (e).
Type: USA, Connecticut, Windsor, 41°51’02”N 72°39’43”W, elevation 38 m, from wood
of Japanese maple (Acer palmatum ‘Thunb., Sapindaceae), 11 June 2013, De-Wei Li
(Holotype, BPI 893194).
EryMOLoey: epithet is named after the genus Acer on which the fungus was collected.
COLONIES on natural substrate thinly diffuse, dark brown. Mycelium partly
immersed and partly superficial, sparse, composed of colorless to pale
brown, irregularly branched, smooth, septate hyphae. CONIDIOGENOUS CELLS
monoblastic, small, 1-celled, smooth, ellipsoid, pale brown, 3 x 2.5 um. CONIDIA
apical, solitary, dry, pleomorphic, of four different forms (as categorized by
Constantinescu & Samson, 1982): form (a) cylindrical, oblong, or clavate,
726 ... Li & Ye
straight or slightly curved, smooth, thick-walled, light brown, (1-)2-septate,
the apical septum covered with a dark band 2.1-4.3 um thick, (11.5-)13.5-16
(-16.6) x (3.5-)4-5.5(-6) um (mean = 14.5 + 1.2 x 5 + 0.6 um, n = 33),
Q = 2.2-4.1 (mean = 3.1), with rounded ends; form (b) oblong or ellipsoidal,
smooth, pale brown, thick-walled, 6.5-11 x 3.5-4.5 um, 1-septate at the middle
or off center; form (d) clavate (incomplete), 17.5-19 x 6-7.5 um, pluriseptate
(often 4-septate), central cells pale brown to brown, with the middle transverse
septum covered by a dark 2-2.5 um wide band; smooth; end cells subhyaline
to light brown, with truncate base; form (f) globose, subglobose, or ellipsoidal,
pale brown to brown, unicellular, smooth, thick-walled, (4—-)4.5-6(-6.5) x
4-4.5(-5) um (mean = 5.1 + 0.7 x 4.2 + 0.3 um, n= 12).
Note: There is a pore visible in the center of each septum and at the basal end of
a conidium. The conidial forms used in this paper follow the six conidial forms
categorized by Constantinescu & Samson (1982). In Triadelphia acericola,
conidia of form (a) predominate, comprising over 90% of all conidia, with the
three other forms (b, d, f) constituting <10%.
Very few intact conidiogenous cells were observed. It appeared that these
conidiogenous cells collapsed once conidia became mature. Two apical cells of
some mature conidia collapsed and even disintegrated, leaving only the basal
cell. No intact form (d) conidia were present. Thus, the number of septa and
size cited for form (d) conidia are approximations.
Conidiogenous cells were also not found in T. inquinans, presumably due to
collapse. Hughes & Pirozynski (1973) and Ellis (1976) did not observe attached
conidia or conidiogenous cells in the specimen of T. inquinans they studied.
Hughes & Pirozynski (1973) assumed that the conidia of T. inquinans develop
singly from swollen hyphal tips (subglobose conidiogenous cells?) that either
collapse after or rupture during conidial secession.
Triadelphia acericola morphologically resembles T! heterospora, T: inquinans,
and T: diversa in having 2-septate, cylindrical, form (a) conidia. Triadelphia
heterospora differs by its much larger form (a) conidia (16.2-19.0 x 3.5-6.0 um;
Shearer & Crane 1971) and the absence of form (b); T: diversa form (a) conidia
differ in their colorless basal cells (Tzean & Chen 1989); and T. inquinans form
(a) conidia differ by the absence of any dark band covering the apical conidial
septum (Hughes & Pirozynski 1973).
Fic. 1—Triadelphia acericola (holotype, BPI 893194). A. form (a) cylindrical conidia; B, C. form
(b) oblong and ellipsoidal conidia; D. form (d) clavate conidia [arrow indicates collapsed terminal
cell]; E. form (a) conidia with form (b) conidia [arrowed]; FE. form (f) globose conidium; G. form (a)
conidia with form (f) conidium [arrowed]; H. conidiogenous cells. Scale bars = 5 um.
Triadelphia acericola & T. centroseptata spp. nov. ... 727
728 ... Li & Ye
Despite several attempts to isolate this fungus onto cultural media, the
fungus failed to grow on either MEA or CMA, suggesting that the fungus
on the specimen may have lost its viability. Failure to isolate this fungus
prevented us from obtaining DNA sequences and carrying out phylogenetic
studies.
Triadelphia alabamensis (Matsush.) Constant. & Samson,
Mycotaxon 15: 482. 1982 FIG. 2B
= Stemphyliomma alabamensis Matsush., Matsush. Mycol. Mem. 2: 16. 1981.
= Pithomyces alabamensis (Matsush.) P.M. Kirk, Trans. Br. Mycol. Soc. 80(3): 462. 1983.
This species develops two different conidial forms: form (d) fusiform or
ellipsoidal, 6-8-septate, 28-44 x 10.5-16 um; end cells colorless or pale brown,
central cells pale brown to brown, dark band covering over the central septum;
form (e) cylindrical, pale olivaceous, 1-3-septate (mostly 1-septate), smooth,
curved, guttulate, with rounded ends, 14-28 x 4.5-5.5 um.
HABITAT/HOsT: On slime flux from wood of Quercus.
DISTRIBUTION: USA (Alabama).
Pithomyces alabamensis is now accepted as the current name of this species
(MycoBank 2017, Species Fungorum 2017). Although its form (e) conidia do
exhibit the rhexolytic secession of Pithomyces, its pleomorphic nature makes
Triadelphia a better fit.
Triadelphia archontophoenicicola D.W. Li, nom. nov. FIG. 2C
MycoBank MB 813055
= Triadelphia australiensis Joanne E. Taylor, K.D. Hyde & E.B.G. Jones, Fungal
Diversity Res. Ser. 12: 366. 2003, nom. illeg. (non B. Sutton 1989).
EryMo_oey: after the genus of Archontophoenix alexandrae (F. Muell.) H. Wendl. &
Drude (Arecaceae) on which the type was collected.
A replacement name (nom. nov.) is required because T: australiensis Joanne
E. Taylor et al. is an illegitimate later homonym of T. australiensis B. Sutton
(McNeill et al. 2012: Art. 53.1).
This fungus develops five conidial forms and two types of conidiophores.
Type 1 conidiophores are micronematous, cylindrical, flask-shaped,
18-30 x 8-12.4 um, pale to dark brown, 1-3-septate; they bear form (d)
conidia, obclavate, 104-132 x 19-24 um, dark brown to black, abruptly
becoming colorless at the long narrow apex, 5-9 um wide, smooth,
10-13 septa, septa in dark part covered in dark bands. Type 2 conidiophores
are semi-macronematous, cylindrical, 60-190 x 4-6 um, dark red-brown,
4—8-septate; they develop form (a) conidia, cylindrical to fusiform, 24-35 x
Triadelphia acericola & T. centroseptata spp. nov. ... 729
GYOS)
eae SE ae GS
f
Fic. 2—Conidial forms produced by 7 Triadelphia spp.: a. T: acericola; b. T. alabamensis;
c. T. archontophoenicicola; d. T: australiensis B. Sutton; e. T. centroseptata; f. T. corticola;
g. T. disseminata. Line drawings by Rafael FE. Castafieda-Ruiz.
730 ... Li& Ye
6-9 um, brown, 1—3-septate covered by narrow dark bands, smooth, with a
conspicuous dark hilum, with or without a small mucilaginous appendage
at the acute apex. In addition, conidial forms (b), (c), and (g) are associated
with both conidiophore types: form (b) conidia, clavate, 24-26 x 8-11
um, brown, 1-3-septate covered by dark bands, smooth, guttulate; form
(c) conidia, acicular with a long colorless apex, 44 x 6 um, pale yellow-
brown, approximately 8-septate, smooth, acute apex, guttulate; and form
(g) conidia, ellipsoidal to cylindrical, 18 x 6 um, pale brown, straight or
curved, smooth, 4-pseudoseptate, obtuse apex and base, guttulate (Taylor
& Hyde 2003).
HABITAT/HOsT: on a dead rachis of Archontophoenix alexandrae.
DISTRIBUTION: Australia.
This is a species with some features that are unique in the genus; it is the
only species that develops two kinds of conidiophores, and it produces
pseudoseptate form (g) conidia that do not correspond with any of the
conidial forms categorized by Constantinescu & Samson (1982). Taylor
and Hyde (2003) considered that these pseudoseptate conidia represented
a new conidial form for the genus when they assigned their new species
to Triadelphia. Further studies on this species are needed to determine its
phylogenetic relationship with other members of Triadelphia and allied
genera.
Triadelphia australiensis B. Sutton, Sydowia 41: 339. 1989 FIG. 2D
This species develops only form (b) conidia, 8.5-10 x 4.5-6 um (Sutton
1989), apical, solitary, dry, ellipsoidal, obovoid to broadly obovoid, smooth,
thick-walled, 1-septate near the base, thickened, some constricted at the
septum.
Hasirats/Hosts: bark, submerged wood, water damaged plywood from a
rink, and dry wall indoors (Li et al. 2013).
DISTRIBUTION: Australia, Canada, Hong Kong, USA.
Triadelphia australiensis is morphologically very similar to T! uniseptata.
The difference is that T. australiensis B. Sutton has smaller conidia than
T. uniseptata, which are larger (12.5-16 x 6.5-10.5 um; Kirk 1983). Ranghoo &
Hyde (1998) identified Trichocladium uniseptatum [= Triadelphia uniseptata] as
the anamorph of their new genus and species, Ascolacicola aquatica Ranghoo
& K.D. Hyde, but their conidial measurements (6.5-10 x 3.8-6.3 um) match
T. australiensis, rather than T’ uniseptata. Moreover, Réblova (2013) designated
this anamorph as merely “Triadelphia-like?
Triadelphia acericola & T. centroseptata spp. nov. ... 731
Triadelphia centroseptata Venkateshw., S.M. Reddy & S.R. Reddy
ex D.W. Li, sp. nov. FIG. 2E
MycoBank MB 823119
“Triadelphia centroseptata” Venkateshw., S.M. Reddy & S.R. Reddy, Indian
Phytopath. 49(4): 340. 1997 [“1996”], nom. inval. (ICN Arts 40.1, 40.6, 40.7).
Type: India, Andhra Pradesh, Warangal, Daharmassagar Village, on dead rachis of Cocos
nucifera L. (Arecaceae), 22 Nov 1989 (Holotype, IMI 300593; isotype, KUMH HM50).
The species was invalidly published by Venkateshwarlu et al. (1997), because
they presented two herbarium accessions (IMI 300593; KUMH HM50) but
failed to nominate either of them as a type, failed to include the word “type” or
an equivalent, and failed to nominate a single herbarium where the holotype
was conserved (McNeill et al. 2012: Art. 40). Here we validate the species by
designating IMI 30059 as its holotype.
One conidial form (a): cylindrical, 12.5-21 x 4-8.5 um, brown, 1-septate,
septum covered in a wide dark band, smooth.
Hasitat/Host: On dead rachis of Cocos nucifera L.
DISTRIBUTION: India
Note—Triadelphia centroseptata is described and illustrated in detail by
Venkateshwarlu et al. (1997). The type material of this species should be closely
examined in the future to determine whether it truly develops only one form of
conidia. Its conidia resemble those of T. inquinans. However, the latter species
produces smaller cylindrical conidia (a) as well as four other conidial forms.
Triadelphia corticola Manohar., N.K. Rao & D.K. Agarwal,
J. Mycopathol. Res. 39: 109. 2001. FIG. 2F
Manoharachary et al. (2001) identified two conidial forms: form (a) as
narrowly cylindrical rounded at the apex, smooth, 18-21 x 4.5-5.2 um,
2-septate, distal septum covered by a wide dark band, basal one narrow, dark
brown; and form (b) as larger, ovoid, smooth, rounded at the apex, 2-septate,
distal septum covered by a wide brownish to black band, 18-21 x 7-8.3 um.
However, based on the type illustration and conidial secession of the fungus,
it appears that these two conidial “forms” represent a single conidial form that
vary in conidial width, both of which we refer to in our key as form (a).
HaBITAt/HOstT: on bark of an unidentified plant.
DISTRIBUTION: India.
Triadelphia corticola is morphologically similar to T: hungarica in conidial size
and shape. But, T: hungarica develops another conidial form with 6-7 septa
(Révay 1987).
732 ... Li& Ye
Triadelphia disseminata Madrid & J. Edathodu, Persoonia 34: 235.2015. Fic. 2g
This is the second species isolated from human beings (Madrid et
al. 2015). It develops three types of conidia in culture: form (a) (sub)
cylindrical with obtuse ends, pale to dark brown, smooth, thick-walled,
with 1 septum situated above the middle (occasionally 1-celled), often
slightly constricted at the septum, 5-11 x 2.5-4 um; form (c) obclavate
to acicular, straight to slightly curved, colorless to pale olivaceous brown,
smooth, thin-walled, 3-4-septate, 43-227 x 3-4 um, with a truncate base
and a cylindrical body gradually tapering into a long beak with an obtuse
end; and form (e) reniform to allantoid, colorless to light brown, smooth,
thin-walled, 1-celled, with obtuse ends, 5.5-11.5 x 2-3.5 um (Madrid et
al. 2015).
Hasitat/Host: from a disseminated infection in an immuno-
compromised patient.
DISTRIBUTION: Saudi Arabia.
Triadelphia disseminata is morphologically similar to T: pulvinata The
conidial form (c) of T. disseminata are 3-4-septate, while in T. pulvinata
5-7 septate (Maggi et al. 1978).
Triadelphia diversa Tzean & J.L. Chen, Mycologia 81: 630. 1989. FIG. 34
This species develops five conidial forms: form (a), cylindrical, straight or
slightly curved, 13.7-24 x 4-8 um, 1-2-septate; septa covered with wide dark
bands; wall smooth; apical and central cells brown, basal cell colorless or pale
brown; form (b), broadly clavate, 11.5-15.3 x 6.3-8 um, with one transverse
septum near the base, covered with a dark band; smooth; apical cell brown,
basal cell colorless or pale brown, with truncate base; form (d), obclavate,
4-6-septate, 15-26 x 6-7 um; end cells colorless to pale brown, acicular, basal
cells subhyaline to pale brown, truncate, central cells pale brown to brown,
often with dark bands at the septa; form (e), allantoid or reniform, colorless
or pale brown, 2 celled, smooth, thin-walled, 8-16 x 3-5 um; and form (f),
obovate, pale brown, 1-celled, smooth, 5.9-9.8 x 4.3-5.6 um with a truncate
base (Tzean & Chen 1989).
HaBITaT/nHOstT: rotten fallen stem of angiosperm.
DISTRIBUTION: Taiwan.
Triadelphia diversa is morphologically similar to T: acericola, T: heterospora,
and T! inquinans. Triadelphia diversa does not develop conidial form (c);
T. acericola does not have conidial forms (c) and (e); T: heterospora has no
Triadelphia acericola & T. centroseptata spp. nov. ... 733
Fic. 3— Conidial forms produced by 6 Triadelphia spp.: a. T: diversa; b. T: heterospora;
c. I: hungarica; d. T: inquinans; e. T: loudetiae; f. T. morgoensis. Line drawings a-e by Rafael F.
Castaneda-Ruiz; line drawing f from Révay (1993), redrawn by the authors.
734 ... Li& Ye
conidial forms (b) and (c) (Shearer & Crane 1971) and T. inquinans, no
conidial form (e) (Hughes & Pirozynski 1973).
Triadelphia heterospora Shearer & J.L. Crane, Mycologia 63: 247. 1971. FIG. 3B
This is the type species of the genus Triadelphia. Shearer & Crane (1971)
originally described T: heterospora as having two conidial forms: form (a),
cylindrical, 2-septate, rounded at the apex, rounded to slightly truncate at base,
brown, smooth, thick-walled, (14.7—)16.2-19.0(-21.2) x 3.5-6.0 um, the apical
septum covered by a black band and form (d), broad ellipsoid, 4-7-septate,
17.6-23.5 x 8.2-10.6 um, end cells colorless to subhyaline, central cells
subhyaline, brown to very dark brown often with black bands at the septa. After
examination of the type material, Constantinescu & Samson (1982) described
two additional conidial forms for this species: form (e), allantoid or reniform,
colorless or pale brown, 0-1 septate, smooth, thin-walled, rounded at both
ends, 8.5-15 x 3-5 um and form (f), obovate to ellipsoidal, pale brown, 1-celled,
smooth, thin-walled, 6-8 x 4-5 um. However, their illustrations clearly showed
that form (f) conidia were thick-walled. Shearer & Crane (1971) included “form
(e)” conidia among cylindrical form (a) conidia in their illustration; it appears
that they considered “form (e)” conidia to be merely immature cylindrical form
(a) conidia.
Hasirats/Hosts: balsa wood block submerged in fresh water, pine block
submerged in sea water in the air, rain water, litter of Mentha aquatic L. and
Mentha xrotundifolia (L.) Huds. from wetland, and debris from a tree-hollow
(Czeczuga et al. 2007, Goénczél & Révay 2004, Révay 1987). This is an aquatic
fungus, but it has also been found in the air of an orchid greenhouse (Magyar
et al. 2011), and its cylindrical conidia were found in the air indoors in the USA
(pers. obs.).
DISTRIBUTION: Cuba, Hong Kong, Hungary, India, Jamaica, Pakistan,
Poland, USA.
Triadelphia heterospora is morphologically similar to T! acericola, T. inquinans,
and T. diversa. Triadelphia heterospora does not develop conidial forms (b)
and (c) (Shearer & Crane 1971); T: acericola does not have conidial forms (c)
and (e); and T: inquinans, no conidial form (e) (Hughes & Pirozynski 1973);
T. diversa does not develop conidial form (c) (Tzean & Chen 1989).
Triadelphia hungarica Révay, Acta Bot. Hung. 33: 68. 1987. FIG. 3C
This species develops two conidial forms: form (a), clavate or slightly
cylindrical, dark brown, 2-septate with apical septum covered by a dark band,
Triadelphia acericola & T. centroseptata spp. nov. ... 735
17.6-20.8 x 6.4-7 um and form (b), fusiform-ellipsoidal, straight or slightly
curved, 6-7-septate, with a dark band in the middle, truncate at the base, 32-36
x 8-10 um, central cells pale brown to dark brown (Révay 1987).
HABITAT/HOsT: on debris of ?Quercus.
DISTRIBUTION: Hungary
Triadelphia inquinans (Sacc.) S. Hughes & Piroz.,
Can.JJ. Bot, 5022524.:.1973)[" 1972"). FIG. 3D
= Dicoccum inquinans Sacc., Michelia 1(2): 264. 1878.
This species develops five conidial forms. Saccardo (1878) originally
described only one type of conidia: form (b), cylindrical-clavate, 16-20 x
4-5 um, l-septate at middle, with rounded apex and truncate base. Hughes &
Pirozynski (1973) and Ellis (1976) described form (b) and form (d) conidia.
Hughes & Pirozynski (1973) re-described this species based on their study
of three specimens including the type. They described two conidial forms:
form (b) conidia, shorter than those of the type, clavate, with thin-walled frill
(part of conidiogenous cell?), 12-18 x 4-6 um and form (d) conidia, broadly
ellipsoidal, usually curved, 15-18 x 8-9 um, possibly 5-septate (incomplete),
the two middle cells thick-walled, dark brown with the median septum covered
by a dark chestnut-brown band, the penultimate cells paler, and the end cells
colorless to subhyaline.
Constantinescu & Samson (1982) studied the type material and described
three additional conidial forms: form (a), cylindrical, (1-)2-septate, pale brown,
smooth, 12-16 x 3-4 um; form (c), obclavate, 3-5-septate, yellowing brown,
smooth, thin-walled, tip not observed, the base truncate, 13-23 (incomplete)
x 4-5 um; and form (f), obovate to large ellipsoidal, pale brown, 1-celled, pale
brown, smooth, thin-walled, 4-5 x 2.5-3 um. However, the illustration of
Constantinescu & Samson (1982) depict form (f) conidia as thick-, not thin-,
walled; and form (d) conidia as larger (17-24 x 7-9 um, [incomplete]) than
those of Hughes & Pirozynski (1973).
Hasirat/Host: decayed inner bark of Populus nigra var. italica Minchh.,
Populus sp.
DISTRIBUTION: Italy.
Triadelphia loudetiae Maggi, Bartoli & Rambelli,
Trans. Br. Mycol. Soc. 71: 150. 1978. FIG. 3E
Maggi et al. (1978) described only the predominant form (a) conidia:
cylindrical, dark brown, 2-septate, rounded at both ends, smooth, a basal pore
visible, 12-18 x 4.5-5 pm. Constantinescu & Samson (1982) included two
736 ... Li& Ye
additional forms: form (c), obclavate, yellowish brown, sometimes slightly
constricted at septa, smooth, thin-walled, 5-6-septate, 30-105 x 4-5 um,
gradually tapering into a long colorless beak and form (e), allantoid, reniform
or nearly cylindrical, colorless or subhyaline, 0-1-septate, smooth, thin-walled,
PAZ SX 2535 mM:
Hasitat/nHost: from rhizosphere of Loudetia simplex (Nees) C.E. Hubb.
DISTRIBUTION: Ivory Coast.
Triadelphia morgoensis Révay, Stud. Bot. Hung. 23: 63. 1993 [“1992”]. FIG. 3F
This species develops three conidial forms: form (a), cylindrical, straight or
slightly curved, 2-septate, thick-walled, with a rounded apex and a truncate
base, 16-25.6 x 3.2-4.8 um; form (c), obclavate, subhyaline, 5-7-septate, base
truncate, gradually tapering into a long beak, 40-57.6 x 3.2-4 um; and form
(e), allantoid or reniform, colorless, one-celled, smooth, thin-walled, 4.8-6 x
2-3 um (Révay 1993).
Hasitat/Host: on rotten wood.
DISTRIBUTION: Hungary.
Triadelphia morgoensis differs from T. diversa by the absence of conidial forms
(b), (d), and (f), by its narrower form (a) conidia having wide dark bands
covering their septa (cf. T: diversa form (a) conidia, 13.7-24 x 4-8 um, without
dark bands), and by its 0-septate form (e) conidia (cf. 1-septate in T’ diversa).
Triadelphia pulvinata Maggi, Bartoli & Rambelli,
Trans. Br. Mycol. Soc. 71: 150. 1978. FIG. 4A
Maggi et al. (1978) described the predominant form (a) conidia as solitary,
cylindrical, deep brown, 1-septate, rounded at both ends, with distinct and
smooth wall, basal pore not easily seen, 11.5 x 4 pm. Constantinescu & Samson
(1982) described all three conidial forms for this species: form (a), cylindrical,
pale brown, 1-septate, rounded at both ends, smooth, wall 0.3 um thick, 9-12
x 3-3.4 um; form (c), acicular to obclavate, colorless, slightly curved, smooth,
thin-walled, 5-7 septate, 40-160 x 3-4 um, gradually tapering into a long tail
with rounded apex and truncate base; and form (e), allantoid or reniform,
colorless or pale brown, 1-celled, smooth, thin-walled, rounded at both ends,
6-9.5 x 2-3 um. Some conidia are mostly clavate, asymmetrical, dark brown,
larger, 9-13 x 4.5-6 um; Constantinescu & Samson (1982) considered these to
represent over-mature form (e) conidia.
Hasirats/Hosts: from rhizosphere of Loudetia simplex, dead rachides of
Phoenix canariensis Chabaud (von Arx 1985), and a human eyelid infection.
Triadelphia acericola & T. centroseptata spp. nov. ... 737
c e f
Fic. 4— Conidial forms produced by 6 Triadelphia spp.: a. T. pulvinata; b. T: queenslandica;
c. TI. romanica; d. T: stilboidea; e. T: synnematofera; f. T: uniseptata. Line drawings by Rafael F.
Castaneda-Ruiz.
738 ... Li& Ye
Triadelphia pulvinata is a soil hyphomycete that was reported to cause
eczematoid, scaly, grey lesions on the skin of both eyelids of a male patient in
Saudi Arabia (Al-Hedaithy 2001).
DISTRIBUTION: Canary Islands, Ivory Coast, Saudi Arabia.
Triadelphia pulvinata was originally described with the conidia and
conidiogenous cells developing on ropes of brown hyphae. This raises questions
of whether these hyphal ropes are premature stages of synnemata or whether
this species is intermediate between synnematous and semi-macronematous
members of the genus. Further studies on this species are necessary.
T. pulvinata is morphologically similar to T. romanica, T. stilboidea, and
T. synnematofera. Both T: stilboidea, and T. synnematofera develop synnemata
(Mercado Sierra & Castaneda 1983, Matsushima 1995), while T. pulvinata and
T. romanica do not. Triadelphia pulvinata does not develop conidial forms
(b) and (d) (Maggi et al. 1978), while T: romanica has no conidial form (d)
(Constantinescu & Samson 1982).
Triadelphia queenslandica Matsush., Matsush. Mycol. Mem. 6: 42.1989. Fic. 4B
This species develops three forms of conidia: form (b), clavate, 1-3-septate,
15-30 x 8-13 um, apical cells; brown, paler towards the basal cell; form (d),
fusiform, apices frequently filiform, conidia without apical portion 30-42 x
12-18 um, 5-6-septate, central cells brown black, paler towards both ends,
apical cell about 400 um long, filiform, 0-septate, colorless, often collapsed at
maturity; and form (e), allantoid to reniform, mostly 1-septate, rarely 2-septate,
16-25 x 5-8 um, pale brown (Matsushima 1989).
HaBITAT/HOsT: on rotten leaf of Musa xsapientum L.
DISTRIBUTION: Australia (Matsushima 1989, Photita et al. 2002)
Triadelphia queenslandica, which is morphologically similar to T. heterospora
and T. hungarica, does not develop conidial forms (a) & (c), while T.
heterospora lacks conidial forms (b) & (c) and T: hungarica does not develop
conidial forms c-f. The conidial form (b) of T: queenslandica does resemble the
chlamydospores produced by aerial mycelia of Trichocladium opacum (Corda)
S. Hughes (Kendrick and Bhatt 1966, Matsushima 1989). However, conidial
forms (d) & (e) of T: queenslandica separate it from Tric. opacum.
Triadelphia romanica Constant. & Samson, Mycotaxon 15: 482. 1982. FIG. 4c
This species develops four forms of conidia: form (a), cylindrical, 0-1-septate,
constricted in the middle, pale brown, smooth, 6.5-14 x 2.5-4 um; form (b),
clavate, brown, smooth, thick-walled (0.5-0.8 tm thick), l-septate, 13-22 x
Triadelphia acericola & T. centroseptata spp. nov. ... 739
5.5-5 um; form (c), obclavate to acicular, colorless, smooth, thin-walled,
2-4-septate, 50-80 x 3-4.5 um, base truncate, gradually tapering into a long
beak of 1-1.5 um wide; and form (e), allantoid to reniform, colorless, 1-celled,
thin-walled, smooth, rounded at both ends, 7-12 x 2-3 um (Constantinescu &
Samson 1982).
Hasirats/Hosts: Isolated from avian eggshell; decayed frond of Palmae
(Matsushima 2001).
DISTRIBUTION: Peru, Romania.
Triadelphia romanica is morphologically similar to T: pulvinata. Triadelphia
romanica does not develop conidial form (d) (Constantinescu & Samson 1982),
while. T: pulvinata has no conidial forms (b) and (d) (Maggi et al. 1978).
Triadelphia stilboidea Mercado & R.F. Castafieda,
Revista Jard. Bot. Nac., Univ. Habana 4(2): 68. 1983. FIG. 4D
This species has synnematous conidiophores, tretic conidiogenous cells, and
two conidial forms; form (a) predominating: cylindrical, sometimes ellipsoidal
or obclavate, olivaceous or brown olivaceous, 1-septate, rarely 0-septate, 7.5-15
x 2.4-4.7 um; and form (?c), seta-like, very long, obclavate, rostrate, 1-celled,
dark brown at the base, becoming paler towards apex, subhyaline at apex,
20-85 x 3.5-4.7 um, tapering to 0.7-1 um at apices. The seta-like conidia of this
species do not fit any forms categorized by Constantinescu & Samson (1982).
Hasirats/Hosts: decayed leaf of unknown palm; dead leaf petiole of
Roystonea regia (Kunth) O.F. Cook.
DISTRIBUTION: Cuba, Puerto Rico.
Révay (1993), who considered T: stilboidea not closely related to T: heterospora
and other species of Triadelphia, excluded it from Triadelphia.
Matsushima (1995) considered that the synnematous Triadelphia stilboidea
has setae similar to those of Trichurus spiralis Hasselbr., not setae-like conidia
as described by Mercado Sierra & Castaneda (1983).
Triadelphia stilboidea and T. synnematofera both develop synnemata
(Mercado Sierra & Castafieda 1983, Matsushima 1995). Triadelphia stilboidea
develops conidial forms (a) and (c) (Mercado Sierra & Castafieda 1983), while
T. synnematofera has conidial form (a) and (e) (Matsushima 1995).
Triadelphia synnematofera Matsush.,
Matsush. Mycol. Mem. 8: 38. 1995, as “synnematfera” FIG. 4E
Synnemata solitary, gregarious, cylindrical, 210-350 um in height. Conidia
of two forms: form (a) cylindric-ellipsoidal, 1-septate, smooth, dark brown,
740 ... Li & Ye
9-17 x 3.5-5 um and form (e) allantoid, reniform or cylindrical, 1-septate,
straight or slightly curved, smooth, colorless, 11-16 x 4-5.5 um (Matsushima
1995).
Hasitat/Host: on decayed petiole of palm
DISTRIBUTION: Peru.
Triadelphia synnematofera is another synnematous species with two conidial
forms. Its form (a) conidia that are 1-septate and have no dark band over the
septum easily distinguish it from T° stilboidea.
Triadelphia uniseptata (Berk. & Broome) P.M. Kirk,
Trans. Br. Mycol. Soc. 80: 464. 1983. FIG. 4F
= Sporidesmium uniseptatum Berk. & Broome, Ann. Mag. Nat. Hist., Ser. 3, 3: 360. 1859.
= Trichocladium uniseptatum (Berk. & Broome) S. Hughes &
Piroz., Can. J. Bot. 50(12): 2526. 1973 [“1972”].
= Dicoccum apiosporum Sacc., Nuovo G. Bot. Ital. 22: 71. 1915 (Hughes & Pirozynski 1972).
= Polyschema bicellulare Shearer, Mycotaxon 14(1): 91. 1982 (Kirk 1983).
This is another species that develops only one conidial form, form (e),
obovoid, 1-septate, smooth, brown, apical cell much larger than the basal
cell. Triadelphia uniseptata is morphologically similar to T. australiensis
B. Sutton, but differs in its larger conidia, 12.5-16 x 6.5-10.5 um (Kirk 1983).
HABITATS/HOSTs: on dead wood, submerged wood, stems, or twigs;
debris from a tree hollow, on bark of Clematis vitalba L., bark of Melicoccus
bijugatus Jacq., dead bark of Quercus rubra L., cones of Pinus halepensis
Mill., rotting wood of Hedycarya arborea J.R. Forst. & G. Forst., stem of
Rubus plicatus Weihe & Nees, on twigs of Platanus occidentalis L., softwood
label in greenhouse, dust from indoor environments, rain water (Berkeley
& Broome 1859, Hughes & Pirozynski 1973, Kirk 1983, Matsushima 1987,
Révay 1987).
DISTRIBUTION: Canada, Cuba, France, Hong Kong, Hungary, India,
Malta, New Zealand, UK, United States.
Key to species of Triadelphia
1. Conidia-only onefornrandnot-synnematous. 9.2 fss.t feat feat feat Lent Stes 2
1. Conidia pleomorphic (2-5 forms) or synnematous ............. 0... cece eee eee 5
PRSONICIA NBEO PALEes ats wine acd ne esis ean Baas ean eee oe se bess Paes MBAS 3
2. Conidia 2-septate, cylindrical or clavate, smooth, 18-21 x 4.5-8.3 um ... T. corticola
3. Conidia 1-septate near the base, thickened, sometimes constricted at the septum,
ellipsoidal, *to-broadly obovoid; smoothie. cc tga sacs ees nee, ces 4
3. Conidia 1-septate at middle, cylindrical, 12.5-21 x 4-8.5um ...... T. centroseptata
Triadelphia acericola & T. centroseptata spp. nov. ... 741
4 Conidia 8 S-lOkK 415-6 pai 645 oe ak es T. australiensis B. Sutton
te Conical 2.5316 X65 500r5 Ue st tad ak Mek Ane i Mel ne T. uniseptata
SiS VMNCMIG A ROSCRE po nats oars atar hee torte pba ara Sakura Sadenne Sake, webetaie, betel, heB ads 6
Dr SV OGOMNALA ADSI Een fie gc nnet tone ager, melee, gna tempts bon ghee oye CoE onegt a eo ee vA
6. Conidia of 2 forms: one cylindrical and 1-septate; the other seta-like ... T! stilboidea
6. Conidia of 1 form: cylindrical and 1-septate ................... T. synnematofera
7. At least one conidial form is broadly obclavate, fusiform ellipsoidal,
with tblacksalicl atthesepiiacre. AR ie BE PrN PR cs Slt as NEY ee PE sate hae abbey 8
Pe SOU ASA DOVES gat A Anal Male Macal ial AN Metal eat a aR tal Ae AN oe A Anak ie 16
8. Conidia fusiform-ellipsoidal, without acerose end cells ............ T: alabamensis
8. Conidia broadly obclavate or fusiform-ellipsoidal with acerose end cells ......... 9
Oey indrical conidiarabsemt at hat Ase nt Mat eal ee BAe 10
9, Cyhindrical conidigipresents. 2x5 2A cry ears eyes eure res Says bg ln eens 12
10. Conidiophores of two types with 5 conidial forms ........ T. archontophoenicicola
TORINOEAS BD OVE seh Pre te ak SrA N Hr wet aah EL eek Yeas ge h ak baal het ong at. ees 11
11. Conidia fusiform-ellipsoidal, 32-36 x 8-10 um ...............004. T: hungarica
11. Conidia fusiform, 30-42 x 12-18 um... 1. eee eee T: queenslandica
12 Allantondortertitorm-comidia absent 3.688 280 a yet hk ot ee eee te a 13
M2 vAllantoid orton. comidiaspresenit. <2 .' a5 sets See oes ob eh histo ahas 14
13. Cylindrical form (a) conidia 2-septate, apical septum covered
with a dark band, 13.4-15.8 x 4.2-5.4um ........... 0... e eee eee T. acericola
13. Cylindrical form (a) conidia (1-)2-septate without dark band,
pale brown, smooth, 12-16 x 3-4um .... ee eee T. inquinans
14; Clavate-conidiaabsemt:..07tseyed ares woes mbt abe’: whee a what'd whose nb-tls ye! 15
14, Clavate conidia present, cylindrical form (a) conidia 13.7-24 x 4-8 um,
1-2-septate; septa covered with dark 2 um bands, with a colorless base
cell T: diversa
15. (Sub)cylindrical form (a) conidia 1-septate, 5-11 x 2.5-4um ...... T. disseminata
15. Cylindrical form (a) conidia 16.2-19.0 x 3.5-6.0 um, 2-septate, the apical septum
coveredeby a Black bands Wn. nu eh lee hadnt adie hadetna Raden Sade eeeok T: heterospora
fo. Cylindrical fori) tonidia- | -septate Acar ies hha gee ee ee ee 8 17
b6zCylindrical form.(ajsconidia-2-septate: i ..8 Aaa Asctak Anat Anes Bt ee ont oie 18
17. Cylindrical conidia 6.5-14 x 2.5-4 tum; clavate conidia present ....... T. romanica
17. Cylindrical conidia 9-12 x 3-3.5 um; clavate conidia absent ......... T. pulvinata
18. Cylindrical conidia brownish, 9-19 x 3-5 um;
reniform-conidig: l2septate. .1 5.465 .ed-axyeges geo ee ees eee eyes T. loudetiae
18. Cells of the cylindrical conidia differently pigmented, 16-25.6 x 3.2-4.8 um;
reniformi<conitdial-septatec 0080.8 Ae ne ett Ark eet ee es T. morgoensis
742 ... Li & Ye
Discussion
Ranghoo & Hyde (1998) reported that Ascolacicola aquatica had T: uniseptata
as its anamorphic state; however, the conidial size that they reported (6.5-10
x 3.8-6.3 um) corresponds with T. australiensis B. Sutton rather than
T. uniseptata, and Réblova (2013) designated the anamorph as merely
“Triadelphia-like”. Platytrachelon abietis (Réblova) Réblova also develops two
forms of anamorphs morphologically similar to the Triadelphia-like anamorph
of Ascolacicola aquatica (Réblova 2013). Réblova & Seifert analyzed nc28S
sequences of Triadelphia uniseptata from Canada and Platytrachelon abietis; no
close relationship was confirmed, and the molecular data placed T. uniseptata
in the Hypocreomycetidae, Sordariomycetes (Réblova & Seifert unpubl.).
These results suggest that Triadelphia is polyphyletic and its systematic
placement remains unknown. Further phylogenetic studies are necessary.
Acknowledgments
The authors are very thankful to Dr. Rafael F. Castafieda-Ruiz and Dra. Gabriela
Heredia for their critical review of the manuscript. The authors express their great
gratitude to Dr. Rafael E Castafieda-Ruiz for making the line drawings in Figures 2-4
for us. The authors are appreciative to Dr. Agnes Révay (Hungarian Natural History
Museum, Hungary), Dr. Donat Magyar (National Environmental Health Institute,
Hungary), Zhihe Li, (McGill University, Canada), and Yale University Library for their
assistance in obtaining literature otherwise not available to the authors. The authors
are very grateful to Dr. James A. LaMondia (The Connecticut Agricultural Experiment
Station) for his pre-submission review, Dr. Lorelei Norvell for her editorial review, and
Dr. Shaun Pennycook for his nomenclatural review.
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 745-757
https://doi.org/10.5248/132.745
Glischroderma Fuckel
GREGOIRE L. HENNEBERT
Rue de l’Elevage 32, B 1340 Ottignies-Louvain-la-Neuve, Belgium
CORRESPONDENCE TO: hennebertg@scarlet.be
ABSTRACT—Glischroderma and its type species G. cinctum are redescribed and illustrated
with respect to their original description. Subsequent collections of the species show
significant divergences, possibly distinct species, which should be investigated from fresh
material. American collections referred by Korf to Glischroderma based on the presence of
gel and similar conidiogenesis are shown to be distinct from Glischroderma in a number of
characters, and a 1999 phylogenetic study has related that anamorph to Pachyphlodes pfisteri.
Glischroderma is also distinguished from Ostracoderma by decisive characters.
Keyworps—Chromelosporium, Lycoperdellon, peridium, Pezizales, pore
Introduction
In 1973, when describing the Botrytis and Botrytis-like genera of conidial
fungi, Hennebert included Ostracoderma Fr. and Glischroderma Fuckel;
both supposedly produced dichotomously branched Chromelosporium-
like conidiogenous hyphae producing abundant one-celled simultaneous
conidia on denticles, but contained inside a peridium similar to the
peridium of some gasteromycetes, such as Lycoperdon. ‘The interpretation
of Ostracoderma by Juel (1920) and Hughes (1953, 1958) was corrected by
transferring the non-peridiate species to Chromelosporium Corda, with
Ostracoderma |ostracos = shell] remaining monotypic with the peridiate O.
pulvinatum Fr. originally classified by Fries in the Trichodermataceae, close to
the hyphomycetes. Although no type material is available for Ostracoderma
pulvinatum, Hennebert (1973) regarded the peridiate genus Lycoperdellon
Torrend as a possible synonym. The genus Glischroderma was kept separate
746 ... Hennebert
for taxa with verrucose spores, absent in Lycoperdellon, a character ignored
by both Fries and Fuckel but noted by Rea (1913).
In 1994 Richard P. Korf and his mycology class collected a specimen
in the valley of Lost Lodge in New York State referred to Glischroderma
based on its abundant gel [glischros = viscous] and Chromelosporium-like
conidiogenesis; a study of the collection was initiated shortly thereafter but
unfortunately interrupted for a long time. My study of the Korf collections
preserved in CUP herbarium, here presented with reference to the type
material of Glischroderma cinctum from herb. G, is dedicated to Korf’s
memory.
Materials & methods
This study is based on the examination of herbarium specimens from the herbaria of
the Botanical Garden of Geneve, Switzerland (G); of the Department of Plant Pathology,
Cornell University, Ithaca, NY, USA (CUP); and of the Botanic Garden, Meisse, Belgium
(B). A 1960 Olympus FH microscope with positive low phase contrast objectives was
used for the microscopic examinations.
Taxonomy
Glischroderma Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 34, 1870.
ORIGINAL DESCRIPTION: Peridium hemisphaericum, tenax, persistens, demum [at
length] in centro irregulariter fissum, e floccis tenuissimis contextum, furfuraceo-
villosum, basi mycelio tenuissimo cinctum, sporidiis concervatis, minutissimis,
globosis, floccis destitutis [lacking hyphae] repletum.
Obs. Peridium in statu juvenili siccum, album, contextu fibroso, dein in centro
ampliatum [enlarged, extended], argillaceum, annulo albo e peridio juvenili represso
orto [born limited from the young peridium] circumdatum.
TYPE SPECIES: Glischroderma cinctum Fuckel
Glischroderma cinctum Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 35,
1870 PLATES 1, 2, 3B,C, 4B, 5A
ORIGINAL DESCRIPTION: 1. G. cinctum Fuck. Eh. 162. —- Peridio hemisphaerico, usque
ad semiunciam diametro transversali, argillaceo, cum annulo albo fibroso cincto, ore
destituto [lacking a pore], sed demum irregulariter in centro fisso, sporidiis globosis,
uniguttulatis, ca. 4 Mikr. diam., argillaceo-rubellis. Tab. I. Fig. 18 a. Fung. Nat.magn. b.
Sporid. [see PLATE 2A]
Ad verlassenen Kohlerstellen an Kohlenstiickchen, sehr selten, im Herbst. Unweit
der Arnsbacher Briicke, im Winkler Wald.
EryMoLoey; Greek glischros = Latin tenax = viscous, sticky, adherent, gripping.
CONIDIOMA peridiate, hemispherical, pulvinate, not constricted at the
base when mature, <9 mm diam. as dried, larger when fresh and mature
Glischroderma Fuckel ... 747
(‘ampliatum”), arising from a limited (“represso”) subiculum adherent to the
charcoal substrate.
SUBICULUM white, giving rise to the young fruitbody (Fuckel) and exceeding
by 1-2 mm the rind of the mature peridium, made of a network of hyaline
septate branched hyphae, 3.5-5.5 um diam., here and there up to 12 um diam.
PERIDIUM persistent but fragile and easily fissured, with no central pore
(“ore destituto”), ocher-grey to argillaceous brown in color as dried, externally
comprising densely interwoven sinuous, septate, hyaline hyphae, and internally
comprising a lacunar network of right- or broadly angled branched septate
hyaline hyphae, 3.5-5.5 um diam. Peridium at first dry when young (Fuckel),
becoming later sticky (“tenax”) or glutinous. According to Fuckel, the peridium
dehisces through an irregular split at the center (“irregulariter in centro fisso”)
with no central pore observed.
CONIDIOGENOUS HYPHAE internal, hyaline, septate, 2-6 um diam., seen as
small collapsed pieces on the dried material. The agglutination of the conidia
around pieces of hyphae (a few bearing denticles) on the microscopic slide
and the regular size of the conidia suggest a conidiogenesis similar to that of
Chromelosporium on irregular repeatedly bifurcate, coralloid, branched hyphae
that produce very abundant conidia borne more or less simultaneously on
minute denticles remaining visible all along the hypha, after which the hyphae
collapse and almost disappear (“floccis destitutis”).
ConipiA holoblastic, globose, 3.5-4.2 um diam., hyaline, pale yellow cream
in mass, often sticking together as initially produced, the wall 0.5 um thick and
minutely verrucose.
HOLOTYPE SPECIMEN EXAMINED: Fuckel, Fungi rhenani 162, 1763 labelled by Fuckel:
«N°162 Nassau’s FLora - Glischroderma cinctum Fuckel - Natiirl. Standort: ad carboner
- Fundort: Oestrika Wald - Erscheint: Aut. (scripsit Fuckel) - LEopOLD FUCKEL» [PLATE
1A]. On an old fireplace on debris of charcoal, very rare, in autumn, near the bridge over
the Arnsbach, in the Winkler Forest, (near Oestrich, Germany). Overlabeled in 1894
when incorporated into the Herbier Barbey-Boissier «Ostracoderma pulvinatum Fr (?)
Peridium in stadio juvenili siccum, album, contextu fibrilloso, dein in centro ampliatum,
tenuissimum, glabrum, argillaceum, cum annulo albo, e peridio juvenili represso orto,
circumdatum. In sylvarum locis adustis, rarissime. Autumno. In sylva Hostrichiensi.»
[PLATE 1c], a conflation of Fuckel’s separate published generic and specific descriptions
of Glischroderma cinctum (in herb. G). The holotype specimen consists of fifty fruit
bodies from 1.5 to 9 mm in diam., glued on a piece of cardboard from loose original
material in charcoal debris and recently overwritten «Herbier (G), Fungi rhenani 162,
Ostracoderma pulvinatum Fr.» [PLATE 1B]. A small envelope containing charcoal debris
and small pieces of the fungus is included.
CoMMENTS— The schema by Hennebert (1973, fig. 10) illustrating the fruit
body of Glischroderma cinctum is misleading in that it reflects the more or
748 ... Hennebert
y ‘ Mee
WASSAU’S F! FLORA. ;
f AP Be pA ar V4
y
Natirl. Standort: #< er hans, ~
Pendort: Coote Yok’
“Brseheint: fc f
Leopold Fuckel.
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a
PLATE 1. A-C. Glischroderma cinctum type material (in herb. G).
Glischroderma Fuckel ... 749
PLATE 2. Glischroderma cinctum: A. Fuckel (1870, Plate 1, fig. 18): a. the fruit body, b. the spores;
B-D fruit bodies of the type material, two in side view (showing the flattened position on a cushion
of white mycelium on the black charcoal substrate) and five in top view (showing some extension
of the white subiculum and absence of a pore in the peridium).
750 ... Hennebert
less pedunculate fruitbody as drawn by Malencon (1964, fig. 1.D) for a more
mature fruitbody instead of the “broadly sessile” fruit body sitting on a limited
white hyphal subiculum noted by Fuckel (1870) and Rea (1913).
Another divergence from Fuckel in Hennebert’s (1973) schema is the
presence of a “well defined central pore” based on the Rea (1913) and Malencon
(1964) interpretations of Glischroderma cinctum; Fuckel denied the presence of
a pore.
Glischroderma aff. cinctum PLATES 3A, 5B
Differs from Glischroderma cinctum by: (1) the dehiscence developing from a central
darker circular zone, scarcely conspicuous, that delimits a star-shape splitting of the
peridial wall to form the lips of a central pore; (2) the hyphal structure of the peridium
seen in siccum lacking the compacted hyphal surface observed in the type of G. cinctum
(PLATE 3B).
PERIDIUM made of a lacunar (crisscross) network of hyphae similar to the
peridium internal context of the G. cinctum type specimen. SUBICULUM
hyphae 3.5-6 um diam., septate, loosely interwoven, branched at broad
or right angles, sometimes at opposite sides and enlarged up to 12 um at
branching. CONIDIOGENOUS HYPHAE are 6-8 um diam., branching more
or less dichotomously or coralloid, producing conidia on conspicuous
denticles all along their side. A Chromelosporium-like conidiogenesis
is deduced of the typical branched agglutination of conidia on pieces of
collapsed the hyphae. Conipia are holoblastic, globose, 3.5-—4.2 um diam.,
the walls thin and minutely verrucose showing 4 to 6 warts in cross section,
SPECIMEN EXAMINED: BELGIUM, St. HuBErtTus Forest, Baconfoy: as Catastoma
circumscissum Berk. & M.A. on burnt ground between mosses under Abies excelsa [=
Picea abies], Oct. 1912, Herb. E. Bommer & M. Rousseau, in herb. B, redetermined as
Glischroderma cinctum by V. Demoulin, 1966.
CoOMMENTS—Malencon (1964) describes the formation of the central pore as
observed in the above specimen. Fuckel describes his species as “ore destituto,
sed demum irregulariter in centro fisso’, lacking any apical pore but splitting
irregularly in the center. None of the fruit bodies in the G. cinctum type material
shows such a pore or darkened circular center that could become a pore at the
center of the peridium. The stickiness of the peridium described by Fuckel as
‘tenax’ has been observed only by C. Rea on fresh collections in England, but
not by G. Malencon in Morocco. Would that observation or the character itself
be circumstantial?
Glischroderma Fuckel ... 751
pe B Bee C
PLATE 3. A. Glischroderma aff. cinctum (herb. B, coll. M. Rousseau in Belgium) showing central
apical pore; B, C. Glischroderma cinctum (type): hyphal structure of the peridium, the context (B)
and the compact surface (C) not observed in G. aff. cinctum. (Scale bars: B, C = 5 um)
Glischroderma aft. cinctum might representa second species of Glischroderma.
This possibility should be investigated from fresh collections and their DNA
analysis.
752 ... Hennebert
Pachyphlodes pfisteri Tocci, M.E. Smith & Healy, Ascomycete.org. 7(6): 360, 2015.
ANAMORPH (MITOSPORIC STATE) PLATES 4A,C, 5C
MYCELIAL MAT not peridiate, flat, irregularly extended, <3.5 cm diam., not
constricted at the edge, appressed at the substrate, entirely or partially covered
of a translucent glutinous liquid gel.
MyceELium white, a net of loosely interwoven broad hyphae, 5.3-8.5 um,
(often <17.5 um) wide, generally straight, branching at wide or right angles,
septate, often fasciculate (synnematous) up to 20 mutually anastomosing
hyphae, the fascicles of hyphae becoming erect, sporulating along their side,
bending in the superficial gel layer, when drying forming a translucent shiny and
somewhat rose orange pellicle or membrane, persistent and brittle, enclosing
superficial loose and fasciculate hyphae and covering the entire surface or only
a part of the hyphal mat.
CONIDIOGENOUS HYPHAE hyaline, septate, most often arising from the side
of erect fasciculate hyphae (synnemata) branching at right angles repeatedly and
shortly, branches soon bifurcating in a very irregular dichotomous or coralloid
way, branches covered at maturity all along their length with abundant conidia
born more or less simultaneously on minute 1 um long denticles that remain
visible of the hyphal cell after release of the conidia.
CONIDIA (mitospores) holoblastic, globose, hyaline, white to pale yellow
in mass, 4.2-6.5(-8) um diam., with a wall 0.5 um thick and conspicuously
verrucose all over.
Ho.otype: FH (MES-306); reference sequence, GenBank JN102474 (Healy et al. 2015).
SPECIMENS EXAMINED: USA, NEw York STATE: Alpine, at the north end of Lost
Gorge (Hendershot Gulf), (1) CUP 62646, as Glischroderma cinctum Fckl. (in Korf’s
handwritten notes Glischroderma ?cinctum) on decaying wet leaves col. Mycology 319
Class, id. R.P. Korf, 14.1X.1994, (2) CUP 63540, as Glischroderma americanum, on rotten
wood of birch, col. J. Haudenshield, id. RPK, 20.IX.1995, (3) CUP 63647, on wet leaves,
col. RPK, Lizon et al. id. RPK, 19.VIII.1997. Ithaca, Coy Glen, (4) south slope nr Elm
St, CUP 62652, on soil, leaves, enmeshing debris, col. P. Lizon, id. RPK 12.X.1994, (5)
CUP 62653, on mossy log. col. P. Mullin, id. RPK, 12.X.1994 (one fruit body 13x25 mm),
(6) CUP 62654, idem as CUP 62653. Malloryville, Eames Bog, (7) % way up north
slope, CUP 62650, as Glischroderma n. sp., on leaves, col. id. RPK, 5.X.1994. GenBank
sequence of anamorph AF133160 (Norman & Egger 1999), (8) %4 way up north slope,
CUP 62651, as Glischroderma n. sp., on leaves, twigs, pieces of bark and base of ferns,
col. KT. Hodges, id. RPK, 5.X.1994 [PLATE 4A] sequenced AF133160 by Norman &
Egger (1999). (9) CUP 63648, on wet wood, col. RPK, Lizon et al., id. RPK, 19.VHI.1997.
(10) CUP 63695, on duff, col., id. RPK, 16.[X.1998. [All specimens, (8) excepted, were
annotated by Rosanne Healy (8.VIII.2013) as having spores and hyphae similar to
Pachyphlodes (thyselli)-mitospore mat.]
Glischroderma Fuckel ... 753
Piate 4. A. Pachyphlodes pfisteri (CUP 62651): epigeous anamorph of the truffle, coll. K.J. Hodges
& R.P. Korf, NY State, USA; B. Glischroderma cinctum (type): conidia; C. Pachyphlodes pfisteri
(type): conidia. (Scale bars: B, C = 5 um)
CoMMENTS—Pachyphlodes pfisteri differs from Glischroderma cinctum, as Korf
noted, by the natural substrate being not charcoal but rotting leaves, bark, and
754 ... Hennebert
PiateE 5. A Glischroderma cinctum (type): branched hyphae, collapsed conidiogenous hypha with
denticles, and conidia; B Glischroderma aff. cinctum (herb. Rousseau): conidiogenous hyphae and
conidia. C. Pachyphlodes pfisteri (CUP 62646-62651, mitosporic state): conidiogenous synnema,
conidiogenous hyphae, and conidia. (Scale bars = 10 um)
twigs under beech or oak, mosses, or fern. Beyond its microscopic similarity
in Chromelosporium-like conidiogenesis and verrucose conidia, the species
might remind of Glischroderma by the gel encrusting the superficial hyphae
into a translucent membrane suggesting a peridium. That membrane should
not be called a peridium, an ontogenically distinct structure of hyphae born
with and covering the entire fertile body (peri = all around); the ontogeny of
the gel membrane distinguishes it from Glischroderma. For that reason, I do
Glischroderma Fuckel ... 755
not classify the Pachyphlodes pfisteri anamorph in Glischroderma, the name
adopted by Korf (1994) and in CUP.
With its conidiogenous synnemata, P pfisteri appears more closely related to
the European Chromelosporium terreste (Fr.) M.B. Ellis.
Healy et al. (2015) published an illustrated description of the epigeous
mitosporic mats of Pachyphlodes pfisteri. Another collection is illustrated here
(Plate 4A, C).
Cultures, attempted by Korf, of broad hyphae on streptomycin CMA on
22 September 1994, grew rapidly, up to 2.5 cm in 10 days (Korf, handwritten
notes). Conidia showed no sign of germination, as in some Chromelosporium
species. For that reason, Healy et al. (2015) suggested that they might function
as spermatia, germinating only in fertilizing ectomycorrhizal hypogeous
hyphae.
The fresh collection of CUP material submitted by Korf for DNA analysis
and published by Norman & Egger (1999), demonstrated a strong support for
monophyly within Pachyphloeus and the Scabropezia clade in the Pezizaceae.
This support was confirmed by Hansen et al. (2001), Tedersoo et al. (2006) and
Leessoe & Hansen (2007). A refined phylogenetic study by Healy et al. (2013)
of all known and undescribed taxa of Pachyphloeus Tul. & C. Tul., nom. illeg.
[= Pachyphlodes Zobel] revealed the existence of 18 new taxa, two of which
were fully described by Healy et al. (2015) as new species, Pachyphlodes pfisteri
from North America and P nemoralis from Europe.
Discussion
The classification of Glischroderma and Ostracoderma near Chromelosporium
by Hennebert (1973) was based on the Malencon (1964) description of
a Moroccan collection of Glischroderma. That observation on the genus
Glischroderma incited Hennebert, also influenced by Juel (1920), to refer
it to Ostracoderma and to synonymize Lycoperdellon, as no type existed for
Ostracoderma pulvinatum and none of the terms of the original description
suggested such interpretation.
Comparison of the original Latin descriptions (TaBLE 1) clarifies the diagnosis.
The only characters distinguishing Glischroderma from Ostracoderma are the
circular hyphal subiculum and the viscosity of the mature fruit body, which
justify the generic name and specific epithet (cinctum). The stickiness of fresh
material observed by Fuckel was not seen by Fries.
Dried and with no traces of viscosity, the type material of Glischroderma
cinctum could easily be identified as Ostracoderma pulvinatum, as has been
done in herb. G, when no attention is drawn to the white circular subiculum
756 ... Hennebert
TABLE 1. Comparative descriptions of Ostracoderma and Glischroderma
Ostracoderma Fr. 1825 Glischroderma Fuckel 1870
Type: Ostracoderma pulvinatum Fr. 1829 Type: Glischroderma cinctum Fuckel 1870
PERIDIUM
Rotundatum, rotundato-hemisphaericum hemisphaericum
crustaceum, crustosum persistens
— in juvenili siccum, [deinde glutinosum]
= tenax,
glabrum, laeve, minime villosum furfuraceo-villosum, e floccis tenuissimis contextum
fragile, fragillissimum in centro irregulariter fissum
in medio fatiscens, collabens, evanescens dein in centro ampliatum, ore destituto
albidum albidum, dein argillaceum
tenue, tenuissimum, % unc. semi unciam diametro
basi mycelio tenuissime cinctum, cum annulo albo
thallus nullus conspicuus ; : Thome nigh
fribroso cinctum, e peridio juvenili represso orto
SPORIDIA
coacervata, laxa, laxe coacervata coacervata
floccis nullis intertexta floccis destitutis
globosa globosa; uniguttulata
argillacea argillaceo-rubellis
minuta minutissima, ca. 4 Mik. diam.
(apparently denied by Fries’s words “thallus nullus conspicuus”).
The words ‘fatiscens, ‘collabens; ‘evanescens’ suggest that Fries observed the
fungus living. Such observation does not match either any central irregular
splitting in Glischroderma cinctum and in Lycoperdellon torrendii (Bres.)
Torrend or any central star-shaped opening to form a pore in Glischroderma aff.
cinctum. In light of these reflections, Ostracoderma remains a mystery and the
earlier proposed synonymy of Lycoperdellon with Ostracoderma (Hennebert
1973) should be regarded now as tentative and even doubtful.
Acknowledgements
My gratitude is to Richard P. (Dick) Korf for his constant
friendship, guidance, and support during 44 professional
years and later. I met Dick for the first time in Ottawa in
1960. He invited us, Lidwina and me, to his home along the
Lake in Ithaca, in the summer 1961. At his door, when we
arrived, we found the message “Welcome, come in, the house
is yours, we come very soon, Dick.’ Surprising! In September
1969, we discussed days and nights the nomenclature of the
pleomorphic fungi for the Kananaskis First Conference,
about my distinction between botanical and anatomical
nomenclature. I was greatly honored when, as Fulbright
Scholar in 1972-73, he chose to stay with us at the University of Louvain. From our joint
wishes exchanged between September 2 and December 10, 1973 in Ithaca, MycoTAXON
Glischroderma Fuckel ... 757
was planned and then born in September 1974. And so our working friendship
continued. Many thanks to You, Dick!
I am also grateful to Dr Philippe Clerc of the Herbarium (G), and to herbarium
curators Dr Scott LaGreca (CUP) and Dr A. Bogaerts (B). My sincere thanks are for
Prof. Dr Rosanne Healy (University of Florida) and Prof. Dr Kathie Hodges (Cornell
University) for their collaboration and the scientific revision of the manuscript. The help
of Dr Cony Decock of MUCL, and of Dr Lorelei Norvell and Dr Shaun Pennycook of
MycotTaxon is also warmly acknowledged.
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Fries EM. 1829. Systema Mycologicum 3(1): 1-259.
Fuckel L. 1870. Symbolae Mycologicae. Beitrage zur Kenntniss der rheinischen Pilze. Jahrbiicher
des Nassauischen Vereins fiir Naturkunde 23-24: 1-459.
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Healy RA, Hobart C, Tocci GE, Bona L, Merényi Z, Paz Conde A, Smith ME. 2015. Fun with the
discomycetes: revisiting collections of Korf’s anamorphic Pezizales and Thaxter’s New England
truffles leads to a connection between forms and the description of two new truffle species:
Pachyphlodes pfisteri and P. nemoralis. Ascomycetes.org. 7(6): 357-366.
Hennebert GL. 1973. Botrytis and Botrytis-like genera. Persoonia 7(2): 183-204.
Hughes SJ. 1953. Conidiophores, conidia, and classification. Canadian Journal of Botany 31(5):
577-659. https://doi.org/10.1139/b53-046
Hughes SJ. 1958. Revisiones Hyphomycetum aliquot cum appendice de nominibus rejiciendis.
Canadian Journal of Botany 36(6): 727-836. https://doi.org/10.1139/b58-067
Juel. HO. 1920. Uber Hyphelia und Ostracoderma, zwei von Fries aufgestellte Pilzgattungen. Svensk
Botanisk Tidskrift 14: 212-222.
Leessoe T, Hansen K. 2007. Truffle trouble: what happened to the Tuberales? Mycological Research
111: 1075-1099. https://doi.org/10.1016/j.mycres.2007.08.004
Malencon G. 1964. Le Glischroderma cinctum Fuck., sa structure et ses affinités. Bulletin de la
Société Mycologique de France 80(2): 197-211.
Norman JE, Egger KN. 1999. Molecular phylogenetic analysis of Peziza and related genera.
Mycologia 91(5): 820-829. https://doi.org/10.2307/3761535
Rea C. 1913 (“1912”). Glischroderma cinctum Fckl. Transactions of the British Mycological Society
4(1): 64-65, plate 2 [lower]. https://doi.org/10.1016/S0007-1536(12)80009-X
Tedersoo L, Hansen K, Perry BA, Kjoller R. 2006. Molecular and morphological diversity of
pezizalean ectomycorrhiza. New Phytologist 170(3): 581-596.
https://doi.org/10.1111/j.1469-8137.2006.01678.x
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 759-766
https://doi.org/10.5248/132.759
Ellismarsporium gen. nov. and Stanhughesiella gen. nov.
to accommodate atypical Helminthosporium and
Corynesporella species
RAFAEL F. CASTANEDA-Ru1z'*, DE-WEI L1I*“, XtU-GUO ZHANG‘,
BRYCE KENDRICK®, BEATRIZ RAMOS-GARCIA‘, SIMON PEREZ-MARTINEZ*
& DAYNET SOSA°7
' Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT),
Alejandro de Humboldt, Académico Titular de la Academia de Ciencias de Cuba,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
? The Connecticut Agricultural Experiment Station,
Valley Laboratory, 153 Cook Hill Road, Windsor, CT 06095, USA
> Co-Innovation Center for Sustainable Forestry in Southern China,
Nanjing Forestry University, Nanjing, Jiangsu 210037, China
‘Department of Plant Pathology, Shandong Agricultural University,
Taian, Shandong 271018, China
° Mycologue, 8727 Lochside Drive, Sidney, BC V8L 1M8, Canada
° Universidad Estatal de Milagro (UNEMI), Facultat de Ingenieria,
Milagro, Guayas, Ecuador
” Escuela Superior Politécnica del Litoral, ESPOL, CIBE,
Km 30.5, Via Perimetral, Guayaquil, Ecuador
* CORRESPONDENCE TO: rfcastanedaruiz@gmail.com
ABSTRACT— Two new genera are proposed: Ellismarsporium for species of Helminthosporium
and Corynesporella that have catenate conidia and polytretic conidiogenous cells, and
Stanhughesiella for a species of Helminthosporium with dictyoseptate conidia. Descriptions
and illustrations are provided.
KEY WORDS—asexual ascomycota, hyphomycetes, taxonomy, tropical fungi
Introduction
Helminthosporium Link, typified by H. velutinum Link, is characterized
by macronematous, unbranched, erect conidiophores. The conidiogenous
760 ... Castaftieda-Ruiz & al.
cells are polytretic, integrated, terminal and intercalary, determinate, with the
conidiogenous loci at the apex and laterally below the septa. The conidia are
solitary, obclavate to cylindrical-obclavate, distoseptate, sometimes rostrate,
smooth, subhyaline to olivaceous brown, usually with an evident dark brown
or brown basal scar.
Helminthosporium dictyoseptatum produces dictyoseptate conidia in
polytretic conidiogenous cells (Hughes 1980). For this species, clearly
separated from the Helminthosporium generic concept, we propose the new
genus, Stanhughesiella.
Several other species with the same conidial ontogeny but producing
blastocatenate conidia have been included in Helminthosporium (Bhat & Sutton
1985, Castaneda-Ruiz & Kendrick 1991, Ellis 1961, Matsushima 1975, Sutton
1993), requiring an expanded generic concept. A similar situation is found in
Corynesporella Munjal & H.S. Gill [typified by C. urticae Munjal & HLS. Gill
(Munjal & Gill, 1961)], which is diagnosed by macronematous conidiophores
and monotretic, terminal or discrete conidiogenous cells that produce
solitary or short catenate, distoseptate, subhyaline or pale brown conidia. In
contrast, Corynesporella pinarensis and C. simpliphora are characterized by
unbranched conidiophores and polytretic, integrated, intercalary and terminal
conidiogenous cells that produce catenate distoseptate conidia (Castaneda-
Ruiz 1985, Matsushima 1993)—characters that deviate from the Corynesporella
generic concept. Consequently, we propose the new genus Ellismarsporium to
accommodate these atypical Helminthosporium and Corynesporella species.
Taxonomy
Ellismarsporium R.F. Castafieda & X.G. Zhang, gen. nov.
MycoBank MB 807603
Differs from Helminthosporium by its acropetal, unbranched or branched chains
of conidia and from Corynesporella by unbranched conidiophores and polytretic
conidiogenous cells.
TYPE sPECIES: Helminthosporium hypselodelphyos M.B. Ellis [= Ellismarsporium
hypselodelphyos (M.B. Ellis) R.E. Castafieda & X.G. Zhang]
EryMo_oey: Latin, Ellismar-sporium, named in honor to Martin Beazor Ellis, (United
Kingdom) for his contribution to the study of hyphomycetes + -sporium, referring to
the conidia.
CoLonliEs effuse, hairy, dark. Mycelium mostly immersed. CONIDIOPHORES
distinct, single, unbranched, straight or flexuous, cylindrical or subulate, mid
to very dark brown, smooth or verruculose. CONIDIOGENOUS CELLS polytretic,
integrated, terminal and intercalary, determinate, cylindrical. CONIDIOGENOUS
New genera: Ellismarsporium & Stanhughesiella ... 761
Fic. 1. Ellismarsporium species: conidiogenous cells and conidia. A. E. hypselodelphyos (Ellis
1961); B. E. catenatum (Matsushima 1975); C. E. parvum (Castafeda-Ruiz & Kendrick 1991);
D. E. pinarense (Castaneda-Ruiz 1985). Scale bars: A, C, D = 10 um; B = 20 um. Redrawn by R.E
Castaneda-Ruiz.
762 ... Castafieda-Ruiz & al.
Fic. 2. Ellismarsporium species: conidiogenous cells and conidia. A. E. senseletii (Bhat & Sutton
1985); B. E. simpliphorum (Matsushima 1993); C. E. zombaense (Sutton 1993). Scale bars:
A, B = 10 um; C = 20 um. Redrawn by R.F. Castafieda-Ruiz.
New genera: Ellismarsporium e& Stanhughesiella ... 763
LOCI pores near septa. Conip1a blastocatenulate, acropleurogenous, obclavate,
cylindrical, ellipsoid to oblong, distoseptate, pale brown to brown, smooth or
verruculose, sometimes with a dark brown or black scar at the ends.
Ellismarsporium hypselodelphyos (M.B. Ellis) R.F. Castafieda & X.G. Zhang,
comb. nov. Fic. 1A
MycoBank MB 807604
= Helminthosporium hypselodelphyos M.B. Ellis, Mycol. Pap. 82: 18 (1961).
Ellismarsporium catenatum (Matsush.) R.F. Castafieda & X.G. Zhang,
comb. nov. Fic. 1B
MycoBank MB 807605
= Helminthosporium catenatum Matsush., Icon. Microfung. Matsush. Lect.: 83 (1975).
Ellismarsporium parvum R.F. Castafieda & W.B. Kendr., nom. nov. Fig. 1C
MycoBank MB 807606
= Helminthosporium parvum R.E. Castafieda & W.B. Kendr., Univ.
Waterloo Biol. Ser. 35: 57 (1991), nom. illeg. (non Grove 1886).
Ellismarsporium pinarense (R.F. Castafieda) R.F. Castafieda & X.G. Zhang,
comb. nov. Fic. 1D
MycoBank MB 807607
= Corynesporella pinarensis R.F. Castafieda, Deuteromyc. Cuba, Hyphomyc. 3: 12 (1985).
Ellismarsporium senseletii (Bhat & B. Sutton) R.F. Castafieda & X.G. Zhang,
comb. nov. Fic. 2A
MycoBank MB 807608
= Helminthosporium senseletii Bhat & B. Sutton, Trans. Brit. Mycol. Soc. 85: 119 (1985).
Ellismarsporium simpliphorum (Matsush.) R.F. Castafieda & X.G. Zhang,
comb. nov. Fic. 2B
MycoBank MB 807609
= Corynesporella simpliphora Matsush., Matsush. Mycol. Mem. 7: 47 (1993).
Ellismarsporium zombaense (B. Sutton) R.F. Castafieda & X.G. Zhang,
comb. nov. Fre. 2€
MycoBank MB 807610
= Helminthosporium zombaense B. Sutton, Mycol. Pap. 167: 32 (1993).
Key to Ellismarsporium species
f OnIdta NOtINOLe than S-GastOSep tate fake A at edctat oat Alaeeat Set ek Bs 2.
LAC onidia more that s-distosepiate ee yal Melk hPa iar pele i daeh pri teeh pits 3
2. Conidia 1-2-distoseptate, cylindrical-ellipsoidal to oblong, slightly constricted
at the middle, smooth, brown, 12-27 x 7-12 um ...............-.. E. parvum
2. Conidia 1-3-distoseptate, fusiform, clavate or oblong, smooth,
pale olivaceous brown, 15-28 x 6.5-8 um .............004. E. hypselodelphyos
764. ... Castafieda-Ruiz & al.
3-Conidia normore than. 6-distoseptate...-.45. 0.55 i621 ws tate eet wns ete ee, a
3.Conidia more that 6=distoseptate’ s+ cess v sews enue een nw OR Oe aS ede bee 6
4, Conidia 1-5-distoseptate, obclavate, smooth, brown to olivaceous brown,
ZS TOPE Seah PDS Beacon ard Ware ne, andes tes tetera conn Pa haere ate 0h E. pinarense
4) Conidianotas above <2 05 h0c bl kuch) hee La eee hse) woe) eevee ween donned erly 5
5. Conidia 0-6-distoseptate, cylindrical to fusiform to obclavate,
slightly constricted at the septa, smooth, mid brown, curved,
OSD SCSI ge. ds aoa c-ngtans api tas-apticne atic extyety a bong erence ® E. senseletii
5. Conidia 2-6-distoseptate, obclavate, curved, smooth, pale olivaceous,
| fears (Obes ci biaete MS UF WR is BRA CR ni A dh By Aes RAE Str SAL ROSE PENCE RL AE E. catenatum
6. Conidia 1-7-distoseptate, obclavate to cylindrical, or long oblong,
curved, pale olivaceous, with brown scar at the ends,
NS OP Armee O QUIT ciate, cues aye cet gettin on gs i0-6--a ga cece ni alias E. simpliphorum
6. Conidia 3—5(-10)-distoseptate, obclavate, smooth, subhyaline,
with a dark brown basal scar, 26-50(-74) x 7.5-9.5 um ......... E. zombaense
ComMENTs: Helminthosporium velutinum, H. dalbergiae, and H. magnisporum
were found phylogenetically associated with the family Massarinaceae by
Tanaka et al. (2015), but no molecular data were found on the seven proposed
Ellismarsporium species; further molecular study is necessary in order to
establish their phylogeny.
Stanhughesiella R.F. Castaftieda & D.W. Li, gen. nov.
MycoBAank MB 807614
Differs from Helminthosporium by its dictyoseptate conidia.
TYPE SPECIES: Helminthosporium dictyoseptatum S. Hughes
[= Stanhughesiella dictyoseptata (S. Hughes) R.F. Castaneda & D.W. Li]
ErymMo.oey: Latin, Stanhughesiella, described in honor to Stanley J. Hughes
(Canada) for his contribution to the studies of the hyphomycetes.
CoLonliEs effuse, hairy, dark. Mycelium mostly immersed. CONIDIOPHORES
distinct, single, unbranched, straight or flexuous, septate, cylindrical or clavate,
mid to very dark brown, smooth or verruculose. CONIDIOGENOUS CELLS
polytretic, integrated, terminal and intercalary, determinate. Conrp1a solitary,
acropleurogenous, simple, obclavate, cylindrical to ellipsoid, dictyoseptate,
pale brown to dark brown, smooth, sometimes with a dark brown or black scar
at the base.
Stanhughesiella dictyoseptata (S. Hughes) R.E. Castafieda & D.W. Li,
comb. nov. FIG. 3
MycoBank MB 807615
= Helminthosporium dictyoseptatum S. Hughes, New Zealand J. Bot. 18: 69 (1980).
New genera: Ellismarsporium e& Stanhughesiella ... 765
Fic. 3. Stanhughesiella dictyoseptata: conidiogenous cells and conidia (Hughes 1980).
Scale bar = 10 um. Redrawn by R.F. Castafeda-Ruiz.
ComMENTSs: Briansuttonia R.F. Castafieda et al., Embellisia E.G. Simmons,
Gibbago E.G. Simmons, and Ulocladium Preuss (Seifert et al. 2011)
are superficially similar to Stanhughesiella, but those four genera have
766 ... Castafieda-Ruiz & al.
cicatrized conidiogenous loci and some produce conidia in acropetal chains.
Stanhughesiella also resembles Stemphylium Wallr. (Seifert et al. 2011), which
is distinguished by monoblastic conidiogenous cells with a distinct cupulate
cicatrized scar that develops after several enteroblastic percurrent regenerations.
Acknowledgments
We are indebted to Dr. Josiane S. Monteiro and Dr. Gabriela Heredia for critical
review. RFCR and BRG are grateful to Organizacion Superior de Direccién Empresarial
Grupo Agricola (OSDE) from Cuban Ministry of Agriculture and Programa de Salud
Animal y Vegetal (project P131LH003033). Dr. Lorelei L. Norvell’s editorial review and
Dr. Shaun Pennycook’s nomenclature review are greatly appreciated.
Literature cited
Bhat DJ, Sutton BC. 1985. New or interesting hyphomycetes from Ethiopia. Transactions of the
British Mycological Society 85: 107-122. https://doi.org/10.1016/S0007-1536(85)80160-1
Castaneda-Ruiz RE 1985. Deuteromycotina de Cuba, Hyphomycetes. HI. La Habana. 42 p.
Castafieda-Ruiz RE, Kendrick B. 1991. Ninety-nine conidial fungi from Cuba and three from
Canada: II. University of Waterloo Biology Series 35. 132 p.
Ellis MB. 1961. Dematiaceous hyphomycetes III. Mycological papers 82. 55 p.
Hughes S.J. 1980. New Zealand Fungi. 27. New species of Guedea, Hadrosporium, and
Helminthosporium. New Zealand Journal of Botany 2: 65-72.
https://doi.org/10.1080/0028825X.1980.10427233
Matsushima T. 1975. Icones microfungorum a Matsushima lectorum. Kobe. 209 p.
Matsushima, T. 1993. Matsushima Mycological Memoirs 7. 141 p.
Munjal RL, Gill HS. 1961. Corynesporella, a new genus of hyphomycetes. Indian Phytopathology
14: 6-9.
Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS
Biodiversity Series 9. 997 p.
Sutton BC. 1993. Mitosporic fungi from Malawi. Mycological Papers 167. 93 p.
Tanaka K, Hirayama K, Yonezawa H, Sato G, Toriyabe A, Kudo H, Hashimoto A, Matsumura
M, Harada Y, Kurihara Y, Shirouzu T. Hosoya T. 2015. Revision of the Massarineae
(Pleosporales, Dothideomycetes). Studies in Mycology 82: 75-136.
https://doi.org/10.1016/j.simyco.2015.10.002
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 767-773
https://doi.org/10.5248/132.767
Endophragmiella jiulingensis sp. nov.
and two new records from China
Hao-Hua Lr, Kat ZHANG’, CHUN-LING YANG’,
JI-WEN XIA* & XIU-GUO ZHANG'*
' Shandong Provincial Key Laboratory for Biology of Vegetable Diseases & Insect Pests,
Department of Plant Pathology, Shandong Agricultural University,
Taian, 271018, China
? Department of Landscaping, Shandong Yingcai University,
Jinan, 250104, China
* CORRESPONDENCE TO: sdau613@163.com, zhxg@sdau.edu.cn
ABSTRACT—A new species, collected on dead branches of an unidentified broadleaf tree from
southern China, is described and illustrated. Endophragmiella jiulingensis is distinguished by
branched conidiophores producing cylindrical, smooth, 0-septate, 6-8 x 4.5-5.3 um conidia.
Botryomonilia scheeleae and Camposporium ramosum are newly recorded from China.
KEY worDs—anamorphic fungi, taxonomy
Introduction
The genus Endophragmiella was erected by Sutton (1973) for two species,
E. pallescens B. Sutton, the type species, and E. canadensis (Ellis & Everh.)
B. Sutton. Subsequently, the genus was emended by Hughes (1979), who gave a
very detailed account of conidiogenesis and generic concepts. Endophragmiella
is mainly characterized by solitary acrogenous conidia seceding rhexolytically
from monoblastic, integrated, terminal, determinate or percurrently extending
conidiogenous cells (Sutton 1973, Ellis 1976, Hughes 1979, Wu & Zhuang 2005).
At present, over 90 species are accepted in the genus (Ma et al. 2011, Ren et al.
2011, MycoBank 2017). All species within Endophragmiella are distinguished
primarily by morphological differences in conidial characteristics, such as
shape, size, pigmentation, septation, and presence or absence of a rostrum at
the apex (Sutton 1973, Ellis 1976, Hughes 1979, Wu & Zhuang 2005).
768 ... Li & al.
A
O
-8B@O @
=
<=
-@@O @
Fic. 1. Endophragmiella jiulingensis (holotype, HSAUP H7927). A, B. Conidiophores,
conidiogenous cells, and conidia; C. Conidia.
Endophragmiella jiulingensis sp. nov. (China) ... 769
During ongoing surveys of saprobic microfungi from forests of southern
China, three interesting anamorphic species were collected from dead branches.
One fungus represents an undescribed species of Endophragmiella B. Sutton,
while the other two species are Botryomonilia scheeleae and Camposporium
ramosum, both new records for China. The specimens are deposited in
the Herbarium of Department of Plant Pathology, Shandong Agricultural
University, Taian, Shandong, China (HSAUP) and the Mycological Herbarium,
Institute of Microbiology, Chinese Academy of Sciences, Beijing, China
(HMAS).
Endophragmiella jiulingensis H.H. Li & X.G. Zhang, sp. nov. FIG. 1
MycoBAank MB 823127
Differs from Endophragmiella corticola by its smaller, 0-septate conidia.
Type: China, Jiangxi Province: Jiuling Mountain, on dead stems of an unidentified
broadleaf tree, 8 Nov. 2016, C.L. Yang (Holotype, HSAUP H7927; isotype, HMAS
245643).
EryMoLoey: in reference to the type locality.
CoLonliEs on the natural substrate effuse, brown. Mycelium partly superficial,
partly immersed, composed of branched, septate, pale brown to brown,
smooth-walled hyphae. CONIDIOPHORES macronematous, mononematous,
arising terminally and laterally from mycelium, erect, straight or flexuous,
irregular branched, multiseptate, smooth, pale brown to brown, 88-187 x
2.9-7 um. CONIDIOGENOUS CELLS integrated, terminal, percurrent, cylindrical,
tapering to a truncate apex, pale brown, 16-29.5 x 2-3 um. Conidial secession
rhexolytic. Conrp1a solitary, acrogenous, cylindrical, 0-septate, smooth, pale
brown to brown, 6-8 x 4.5-5.5 um.
ComMENTS—Endophragmiella jiulingensis is similar to E. corticola P.M. Kirk in
having branched conidiophores, but E corticola differs by its larger, multiseptate
conidia (16-25 x 5.5-7.5 um, 2-3-septate; Kirk 1982).
Botryomonilia scheeleae Goos & Piroz., Canad. J. Bot. 53: 2928, 1975. FIG. 2
Cotonigzs on the natural substratum effuse, brown, hairy. Mycelium
superficial and immersed, simple or branched, dark yellow-brown hyphae.
CONIDIOPHORE stalk <72 um high and with <30 perforated septa, mostly
7.5-12.5 um diam. but sometimes narrowed near the base or wider towards
the tip; wall thin, roughened by fine to coarse vesicular inclusions, which
are distributed evenly along the entire conidiophore length or limited to (or
coarser along) only a part of it. At the apex the conidiophore dichotomously
770... Li & al.
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7
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.
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Fic. 2. Botryomonilia scheeleae (HSAUP H7857). A. Conidiophores, conidiogenous branches,
and conidia; B. Conidiophore and conidiogenous branches; C. Conidiogenous cells and conidia;
D. Conidia.
Endophragmiella jiulingensis sp. nov. (China) ... 771
branched, producing a dendroid, penicillate head of irregular, finger-like
projections. Conrp1a unicellular, hyaline (pale straw-colored when old), held
together in branched chains only by a thin strand of protoplasm connecting
the septal pores; the terminal conidia small (2.5-3 um diam) and more or less
hemispherical, but the lower conidia progressively enlarging (mostly to 6.5 x
6 um, but up to 8 x 4 um) towards the stalk and doliiform.
SPECIMEN EXAMINED: CHINA, GUANGXI PROVINCE, Longhu Mountain, on decaying
twigs of an unidentified broadleaf tree, 7 Dec. 2015, C.L. Yang (HSAUP H7857).
ComMMENTS—Botryomonilia scheeleae is reported for the first time from China.
Botryomonilia is characterized by single differentiated colonies effused on
black stromatic or sclerotial tissue composed of plectenchyma of dark brown
hyphae. Conidiophores erect, simple or branched, resembling conidiophores
of Botrytis, bearing a head of powdery conidia. Heads at first composed of
branched, penicillate conidiogenous cells; later, these branches producing
solitary or catenate conidia by thallic-arthric disarticulation, resembling
Monilia. Conidia unicellular, doliiform, connected by protoplasmic filaments.
The Chinese specimen closely matches the original description of B. scheeleae,
except that the type material had wider conidiophores (15-20 um; Goos &
Piroz. 1975).
Camposporium ramosum Whitton, McKenzie & K.D. Hyde,
Fungal Diversity 11: 183, 2002. FIG. 3
COLONIES on natural substrate effuse, consisting of individual
conidiophores scattered over the substrate surface, brown. Mycelium
immersed and superficial. CONIDIOPHORES 39-87 um long, 3.5-6 um diam.
towards the base, macronematous, mononematous, solitary, unbranched, erect,
irregularly cylindrical, flexuous, dark brown towards the base, fading to pale
brown towards the apex, smooth, 4-10-septate, thickened walls and septa.
CONIDIOGENOUS CELLS holoblastic, polyblastic, integrated into the apical region
of the conidiophores, pale brown, smooth, denticulate; cylindrical denticles
acting as separating cells, 1-3 denticles per conidiophore. Conip1a 50-55.5
um long, 5-6.5 um diam. at the widest point, solitary, dry, cylindrical, brown
or pale brown, slightly thickened walls and septa, 7-9-septate, apex rounded;
basal cell truncate, often with a persistent portion of the denticle attached; the
apical cell giving rise to a single simple or branched appendage; appendage
hyaline, 1-2-septate, smooth, tapering from the base to the apex, close to the
base typically dividing into 3 (rarely 2) terminal branches, individual branches
11-27 um long.
772... Li & al.
Fic. 3. Camposporium ramosum (HSAUP H7919). A-C. Conidiophores, conidiogenous cells,
and conidia; D. Conidia; E. Conidiophore; F. Conidiophores and conidiogenous cells.
Endophragmiella jiulingensis sp. nov. (China) ... 773
SPECIMEN EXAMINED: CHINA, JIANGXI PROVINCE, Jiuling Mountain, on decaying
twigs of an unidentified broadleaf tree, 2 Nov. 2016, C.L. Yang (HSAUP H7919).
COMMENTS—Camposporium ramosum has been reported from Australia and
USA but has not previously been recorded in China. The genus is characterized
by dematiaceous, simple conidiophores that bear terminal, integrated,
denticulate conidiogenous cells. The conidia are typically cylindrical and
elongate multiseptate, rounded at either or both ends; the apex is either simple
or has one or more cylindrical appendages, and the base typically has a persistent
portion of the denticle attached. The Chinese specimen closely matched the
protologue description of C. ramosum, except that the type specimen had
bigger conidia with more septa (80-112 x 6.4-9.6 um, 8-15-septate; Whitton
et al. 2002).
Acknowledgments
The authors express gratitude to Dr. Rafael E Castafieda-Ruiz and Dr. Jian Ma for
serving as pre-submission reviewers and for their valuable comments and suggestions.
This project was supported by the National Natural Science Foundation of China
(Nos. 31093440, 31230001, 31493010, 31493011, 31200013) and the Ministry of Science
and Technology of the People’s Republic of China (Nos. 2006FY120100).
Literature cited
Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew,
Surrey, England.
Hughes SJ. 1979. Relocation of species of Endophragmia auct. with notes on relevant generic names.
New Zealand Journal of Botany 17: 139-188. https://doi.org/10.1080/0028825X.1979.10426887
Kirk PM. 1982. New or interesting microfungi IV. Dematiaceous hyphomycetes from Devon.
Transactions of the British Mycological Society 78: 55-74.
https://doi.org/10.1016/S0007-1536(82)80077-6
Ma LG, Ma J, Zhang YD, Zhang XG. 2011. Taxonomic studies of Endophragmiella from southern
China. Mycotaxon 117: 279-285. https://doi.org/10.5248/117.279
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Goos RD, Pirozynski KA. 1975. Fungi of Barro Colorado Island: new and interesting hyphomycetes.
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Ren SC, Ma J, Zhang XG. 2011. A new species and new records of Endophragmiella from China.
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Sutton BC. 1973. Hyphomycetes from Manitoba and Saskatchewan, Canada. Mycological Psapers
132 143 p.
Whitton SR, McKenzie EHC, Hyde KD. 2002. Microfungi on the Pandanaceae: two new species of
Camposporium and key to the genus. Fungal Diversity 11: 177-187.
Wu WP, Zhuang WY. 2005. Sporidesmium, Endophragmiella and related genera from China. Fungal
Diversity Research Series 15. 351 p.
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 775-778
https://doi.org/10.5248/132.775
Pseudocercospora lysidices sp. nov.
on Lysidice rhodostegia from China
FENGYAN ZHAI”, YINGLAN Guo’, YINGJIE LIu* & HONGLIAN LI”
"Henan Agricultural University, Zhengzhou 450002, China
’ Postdoctoral Research Base, Henan Institute of Science and Technology,
Xinxiang 453003, China
* Institute of Microbiology, Chinese Academy of Science, Beijing 100101, China
* CORRESPONDENCE TO: honglianli@sina.com
ABSTRACT—Pseudocercospora lysidices on Lysidice rhodostegia (Fabaceae subfam.
Caesalpinioideae) is introduced as a new species, based on Chinese specimens previously
misidentified as “Pseudocercospora lathyri.” The new species is described, illustrated, and
compared with other Pseudocercospora species on caesalpiniaceous hosts. The specimens
examined are deposited in the Herbarium of the Institute of Microbiology, Chinese Academy
of Sciences, Beijing (HMAS).
KEY worDs—anamorphic ascomycete, Cercospora, taxonomy, cercosporoid fungi
Introduction
Cercospora lathyri Dearn. & House was described on Lathyrus japonicus
[= L. martimus] (Fabaceae subfam. Faboideae), forming angular leaf spots,
relatively large stromata (30-60 um diam.), short and narrow conidiophores
(5-20 x 3-4 um), and conidia (40-75 x 2-3.5 um), but without any details
on the conspicuousness of the conidiogenous loci (House 1916). Guo & Liu
(1992), who examined a Chinese collection of a cercosporoid hyphomycete
on Lysidice rhodostegia, considered the morphological characteristics of this
fungus to be rather similar to those of Cercospora lathyri, an assumption that
was based only on descriptions of the latter species and without examining the
type material. They assigned the Chinese material to C. lathyri, and introduced
the new combination Pseudocercospora lathyri (Dearn. & House) Y.L. Guo &
776 ... Zhai & al.
X.J. Liu due to unthickened and undarkened conidiogenous loci and hila of the
conidia. Braun & Mel'nik (1997) re-examined the type specimen of C. lathyri
and found small but conspicuously thickened and darkened conidiogenous
loci and hila and subcylindrical-filiform acicular to narrowly obclavate hyaline
conidia, suggesting that C. lathyri must be maintained in Cercospora s. str. and
that the Pseudocercospora found on Lysidice rhodostegia in China represents a
different species, which we describe here as a new species, Pseudocercospora
lysidices.
Materials & methods
The original collection of the specimen in 1961 is described in the Type designation
section. Small pieces of leaf lesions were cut off and heated in Lactic acid-phenol-
glycerol-distilled water (1:1:2:1) between slides and coverslips over flame to become
transparent. The tissues were examined microscopically without any staining using oil
immersion (bright field and phase contrast) lenses on a standard light Olympus CX 21
microscope. Thirty conidia and other structures were measured using 400 magnification;
the measurement ranges include the extremes given in parentheses. The type specimen
is deposited at the Mycological Herbarium of Academia Sinica, Beijing, China (HMAS)
Taxonomy
Pseudocercospora lysidices Y.L. Guo & EY. Zhai, sp. nov. FIG. 1
MycoBAnk MB 819544
MISAPPLIED NAME: “Pseudocercospora lathyri” sensu Y.L. Guo & X.J. Liu (1992).
Differs from Pseudocercospora guanicensis in its distinct amphigenous leaf spots,
its larger stromata, its shorter and narrower pale olivaceous conidiophores, and its
cylindrical-obclavate, paler, shorter, and much narrower conidia.
Type: China, Guangdong Province, Guangzhou, on living leaves of Lysidice rhodostegia
Hance (Fabaceae subfam. Caesalpinioideae), 7 X1 1961, coll. QM. Ma & XJ. Liu
(Holotype, HMAS 59072).
EryMoLoGcy: referring to the host genus Lysidice.
Leaf spots amphigenous, subcircular, angular to irregular, 2-8 mm diam.,
grayish white in the center, margin dark brown or totally dark brown on
the upper surface, brown to dark brown on the lower surface. Caespituli
amphigenous. Mycelium internal. Stromata globose, dark brown, 25-65
um diam. Conidiophores densely fasciculate, pale olivaceous, uniform in
color, irregular in width, not branched, geniculate in the upper part, straight
to curved, obtusely rounded to conical at the apex, 0-1-septate, thin-
walled, smooth, 6.5-22(-30) x 2.5-4.3 um. Conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells, conidiogenous
loci inconspicuous, unthickened, undarkened. Conidia solitary, cylindrical-
Pseudocercospora lysidices sp. nov. (China) ... 777
Fic. 1. Pseudocercospora lysidices (holotype, HMAS 59072).
Conidiophores and conidia. Scale bar = 40 um.
obclavate, subhyaline to very pale olivaceous, straight to curved, obtuse at
the apex, obconically truncate to almost truncate at the base, inconspicuously
3-7-septate, thin-walled, smooth, 24-86.5 x 2.2-3.5 um.
Notes: There are four other Pseudocercospora spp. described on hosts
belonging to the Fabaceae subfam. Caesalpinioideae: Pseudocercospora
caesalpiniae T.K. Goh & WH. Hsieh, P caesalpiniigena U. Braun,
P. caesalpiniicola U. Braun et al., and P. guanicensis (E. Young) U. Braun &
Crous, but they differ in some respects from the new species.
Pseudocercospora caesalpiniae, although also forming angular to irregular
leaf spots, amphigenous caespituli, and short and narrow conidiophores
(5-15 x 1.5-3 um), differs from P lysidices in its external mycelium with solitary
conidiophores, the lack or production of smaller stromata (<40 um diam.), and
its acicular to narrowly obclavate conidia (Goh & Hsieh 1989).
Pseudocercospora caesalpiniigena |= Cercoseptoria caesalpiniae J.M. Yen et
al.] differs from P. lysidices in lacking stromata and in forming indistinct leaf
spots, longer and broader conidiophores (30-60 x 4-5 um), and cylindrical
conidia (Yen et al. 1982, Braun & Freire 2004).
778 ... Zhai & al.
Pseudocercospora caesalpiniicola [= Phaeoisariopsis caesalpiniae J.M. Yen
et al.] also forms large stromata (40-75 um diam.) but is distinguished from
P. lysidices by indistinct leaf spots, synnematous and hypogenous conidiomata,
and longer and broader conidiophores (20-150 x 3-5 um) and conidia (20-110
x 2.5-4.5 um) (Yen et al. 1982, Hernandez-Gutiérrez & Dianese 2009).
In comparison to P. lysidices, PR guanicensis [= Cercospora guanicensis
E. Young] also forms circular to subcircular leaf spots and small stromata
(30-45 um diam.), but its conidiophores are arranged in dense fascicles,
darker in color (olivaceous brown), and much longer and broader (20-215
x 4-8 um) and its conidia are obclavate to obclavate-cylindrical and longer
and broader (40-105 x 5-7 um; U. Braun, pers. comm.; Chupp 1954; Crous
& Braun 2003).
Acknowledgments
This study was supported by the Project for Fundamental Research on Science and
Technology, Ministry of Science and Technology of China (No. 2013FY110400), the
National Natural Science Foundation of China (No. 31100013) and The Cultivation
Program of Youth Backbone Teachers in Colleges and Universities of Henan Province
(2016GGJS-109). We express our deep appreciation to Prof. Uwe Braun and Mark L.
Gleason for their valuable suggestions, kind help, and their assistance during the course
of submission of this manuscript.
Literature cited
Braun U, Freire FDO. 2004. Some cercosporoid hyphomycetes from Brazil - III. Cryptogamie,
Mycologie 25: 221-244.
Braun U, Melnik VA. 1997. Cercosporoid fungi from Russia and adjacent countries. Trudy
Botanicheskogo Instituta imeni V.L. Komarova (St. Petersburg) 20: 1-130.
Chupp C. 1954 [“1953”]. A monograph of the fungus genus Cercospora. Ithaca, NY. 667 p.
Crous PW, Braun U. 2003. Mycosphaerella and its anamorphs: I. Names published in Cercospora
and Passalora. CBS Biodiversity Series 1. 571 p.
Goh TK, Hsieh WH. 1989. New species of Cercospora and allied genera of Taiwan. Botanical
Bulletin of Academia Sinica 30: 117-132.
Guo YL, Liu XJ. 1993. Study on Pseudocercospora in China II. Mycosystema 11: 125-135.
(in Chinese)
Hernandez-Gutiérrez A, Dianese JC. 2009. New cercosporoid fungi from the Brazilian Cerrado 2.
Species on hosts of the subfamilies Caesalpinioideae, Faboideae and Mimosoideae (Leguminosae
s. lat.). Mycotaxon 107: 1-24. https://doi.org/10.5248/107.1
House HD. 1916. New or interesting species of fungi. III. Bulletin of the New York State Museum
188: 29-59.
Yen JM, Kar AK, Das BK. 1982, Studies on hyphomycetes from West Bengal, India, III. Cercospora
and allied genera of West Bengal, 3. Mycotaxon 16(1): 80-95.
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 779-787
https://doi.org/10.5248/132.779
Bactrodesmiastrum domesticum sp. nov. and
Conioscypha varia from indoor environments
DeE-WEI L1*»?, CHIN S. YANG? & ARIUNAA JALSRAI*
' The Connecticut Agricultural Experiment Station Valley Laboratory,
153 Cook Hill Road, Windsor, CT 06095, USA
? Southern China collaborative Innovation Center of Sustainable Forestry,
Nanjing Forestry University, China
° Prestige EnviroMicrobiology, Inc.,
242 Terrace Boulevard Suite B-1, Voorhees, New Jersey 08043, USA
*EMLab Pe&K, 3000 Lincoln Drive East, Suite A, Marlton, NJ 08053, USA
* CORRESPONDENCE TO: dewei.li@ct.gov
ABSTRACT—Two hyphomycetes were collected from residences: a new species,
Bactrodesmiastrum domesticum, with turbinate or obtriangular conidia and Conioscypha
varia, reported for the first time from an indoor environment.
Key worps—building, dry wall, asexual fungi, water damage, wood
Introduction
Indoor environments provide man-made and unique habitats for fungi.
Building materials, such as drywall and wood, and household materials such
as clothing, fabric, books, paper products, foodstuffs, etc. are good substrates
for fungi whenever water or moisture is available. There have been historical
studies on indoor fungi (e.g., the notorious Stachybotrys chartarum (Ehrenb.)
S. Hughes [= Stachybotrys atrus Corda] collected from an indoor wall in Prague
during the early 19 century; Corda 1837), but fungal diversity in indoor
environments has not been well studied, perhaps due to the misconception
that fungal diversity in indoor environments is simple and straightforward.
However, an increasing number of fungi have been added to the list of indoor
fungi in the last two decades, some new to science and others as first records
780 ... Li, Yang & Jalsrai
for the countries where they were collected. We report here on two unique
hyphomycetes were collected from indoor environments in the USA, one a new
species and the other representing a first occurrence for an indoor environment.
Materials & methods
Two fungi were collected during indoor mold inspections in Florida and
Massachusetts. These fungi were grown on malt extract agar (MEA), corn meal agar
(CMA) and dichloran glycerol agar (DG18) at 25°C. A small scalpel was used to
remove fungal structures directly from the samples or from culture for morphological
observation under a compound microscope. Conidiophores, conidiogenous cells, and
conidia were mounted in 85% lactic acid or lacto-fuchsin (0.1 g acid fuchsin, 100 ml
85% lactic acid; Carmichael 1955) and microscopically observed and measured using an
Olympus BX40 compound microscope equipped with Nomarski differential interference
contrast optics and photographed using an Olympus Microfire digital camera (Goleta,
CA). The fungal structure measurements were statistically analyzed using Microsoft
Office Excel 2013 with 95% confidence interval of means. The results are presented as
ranges of the measurement. Specimens were conserved in the Herbarium, Connecticut
Agricultural Experiment Station, New Haven, CT, USA (NHES).
Taxonomy
Bactrodesmiastrum domesticum D.W. Li & Chin S. Yang, sp. nov. PLATE 1
MycoBank MB 820855
Differs from Bactrodesmiastrum monilioides, B. obovatum, and B. obscurum by its much
larger turbinate conidia and from B. pyriforme by its much wider turbinate conidia.
Type: USA, Florida, Miami Shores, unidentified decayed wood (floor plank) in a
residence, 18 March 2013, Albert Baerren, Prestige 130320-01-001 (Holotype, NHES
L1701).
ETyMOLOoGy: epithet is named after the indoor habitat where this fungus was collected.
CotontEs on the natural substrate effuse, sparse, light brown. Mycelia partly
superficial and partly immersed in the decayed wood. Hyphae branched, septate,
pale brown to brown, smooth, thick-walled, 2.5-3 um diam. CONIDIOPHORES
poorly differentiated or undifferentiated, often reduced to conidiogenous cells,
very rarely developing a septum and separated into two cells, straight, rarely
flexuous, unbranched, solitary. CONIDIOGENOUS CELLS monoblastic, smooth,
thick-walled, pale brown to brown, much darker than the adjacent hyphal
cells to both sides, (4.5-)3.7-7(-8) um (n = 6) long, (2.5-)3.5-5.5 um (n = 6)
diam at the base, determinate, attenuated towards the apex, conical, tubular,
barrel-shaped or (rarely) ovoid, truncate at the apex, occasionally L shaped
or short inverse T shaped, 2-4.5(-5) um (n = 11) diam. CONIDIAL SECESSION
schizolytic Conip1a (2-)3(-4) septate, acrogenous, turbinate or obtriangular
Bactrodesmiastrum domesticum sp. nov. (USA)... 781
>
Cal, -“~.
Me 2.
b c
PLATE 1. Bactrodesmiastrum domesticum (holotype, NHES L1701). a. Conidia and conidiogenous
cell; b, c. Conidia and conidiogenous cells. Scale bars = 10 um.
with a rounded distal side, black, smooth, glistening, solitary, dry, (36.5-)
39.5-43.5(-47) x (30-)33-37(-37.5) um (n = 30), with distal cells darker than
the proximal cell, conico-truncate at the base. SEXUAL STATE unknown.
Notes: Bactrodesmiastrum Hol.-Jech. is characterized by solitary or poorly
ageregated differentiated or undifferentiated conidiophores reduced to a
conidiogenous cell that is monoblastic, brown, single (Holubova-Jechova
1984) and contrasts with Bactrodesmium, which develops sporodochia and
differentiated, colorless to brown, simple or branched conidiophores that
develop mono- or polyblastic conidiogenous cells (Ellis 1971, Holubova-Jechova
782 ... Li, Yang & Jalsrai
1972). Bactrodesmiastrum was typified by B. obscurum Hol.-Jech. (Holubova-
Jechova 1984), which has flask-shaped conidiogenous cells with a truncate apex
and obovoid versicolored conidia with a conico-truncate basal cell. Hernandez-
Restrepo et al. (2013) added two new species, Bactrodesmiastrum pyriforme
M. Hern.-Rest. et al. and B. obovatum (M. Calduch et al.) J. Mena et al.
[= Janetia obovata M. Calduch et al.].
When describing another new species, B. monilioides Hern.-Rest. et al.,
Hernandez-Restrepo et al. (2015) emended the genus by expanding the
conidiophore description from “reduced to a conidiogenous cell which is
monoblastic, brown, single” to “semi-macronematous or macronematous,
mononematous, simple, usually reduced to conidiogenous cells arranged
singly or aggregated in small groups, or conidiophores arising from pulvinate
to subpustulate, scattered and black sporodochial conidiomata’ All four
previously published species were collected or isolated from decaying or
submerged wood or twigs and exclusively in Europe; B. domesticum represents
the first Bactrodesmiastrum species reported from a different habitat as well as
from another part of the world.
Bactrodesmiastrum domesticum can easily be distinguished from B.
monilioides, B. obovatum, and B. obscurum, which have much smaller non-
turbinate conidia: B. monilioides 24-35 x 15-18 um, B. obovatum 22.5-33.5
x 12-15 um, and B. obscurum 24-35 x 11-15 um. (Hernandez-Restrepo et
al. 2013, 2015; Holubova-Jechova 1984). Bactrodesmiastrum domesticum and
B. pyriforme develop conidia of similar lengths, but those of B. pyriforme are
pyriform and narrower (14-28 um; Hernandez-Restrepo et al. 2013).
Bactrodesmiastrum domesticum also resembles Trichocladium achrasporum
(Meyers & R.T. Moore) Shearer & J.L. Crane, T: nypae K.D. Hyde & Goh,
and T. palmae Manohar. et al. in morphology of conidia and conidiogenous
cells. However, T. achrasporum develops sporodochia and produces much
smaller (17-24 um x 10-16) conidia (some with dark bands at the septa) that
are pyriform, not turbinate (Shearer & Crane 1971); T: nypae develops much
smaller (15-20 x 10-13 um) and pyriform to ellipsoidal conidia (Hyde et al.
1999); and T. palmae has finely verrucose conidia (28-36.5 x 18.5-24 um;
Manoharachary et al. 2006).
Both Bactrodesmiumi linderi (J.L. Crane & Shearer) M.E. Palm & E.L. Stewart
and Ba. moenitum (J.L. Crane & Shearer) M.E. Palm & E.L. Stewart resemble
B. domesticum somewhat in conidial morphology. However, these species
clearly differ from B. domesticum: Ba. linderi by its production of sporodochia
and its much smaller pyriform conidia (15-28 x 11-18 um) and Ba. moenitum
Bactrodesmiastrum domesticum sp. nov. (USA) ... 783
ew cy a)
Sf AN
PLATE 2. Conioscypha varia (NHES L1702). a. Conidia and conidiogenous cells; b. Conidiogenous
cell; c. Conidia; d. Conidium developed on hypha. Scale bars = 5 um.
by its production of sporodochia and its much narrower pyriform conidia (31-
48 x 11-27 um; Palm & Stewart 1982).
784 ... Li, Yang & Jalsrai
Hernandez-Restrepo et al. (2013) found that Ascotaiwania persoonii Fallah
et al., B. obovatum, and B. pyriforme formed a monophyletic clade with 99%
support. (The anamorph of A. persoonii is unknown; Fallah et al. 1999). This
clade was sister (with 99% support) to a /Savoryellales clade that included three
different Ascotaiwania species. Similarly, Hernandez-Restrepo et al. (2015,
in an analysis that did not include A. persoonii) found that B. monilioides,
B. obovatum, and B. pyriforme formed a monophyletic clade with 99% support
that was sister (with 95% support) to a /Savoryellales clade that included
four different Ascotaiwania species (including the type species, A. lignicola).
The extensive phylogenetic analysis by Yang et al. (2016) defined a major
lineage (with 100% support), which they proposed as a new family and order
(Fuscosporellaceae, Fuscosporellales); this included B. monilioides, B. obovatum,
and B. pyriforme, as well as A. persoonii, which they recombined as the type of
a new genus Pseudoascotaiwania Jing Yang et al., remote from Ascotaiwania
sensu stricto (Savoryellales).
Our attempts to culture B. domesticum have been unsuccessful. Trichoderma
koningii Oudem., Scopulariopsis brevicaulis (Sacc.) Bainier, and Fusicladium
aromaticum (Ellis & Everh.) K. Schub. & U. Braun [= Cladosporium aromaticum
Ellis & Everh.] co-existed on the same wood sample.
Conioscypha varia Shearer, Mycologia 65: 133 (1973) PLATE 2
= Conioscyphascus varius Réblova & Seifert, Stud. Mycol. 50(1): 101 (2004)
COLONIES growing slowly on MEA and CMA, no growth on DG18;
reaching 21-24 mm diam on MEA and 22-24 mm diam on CMA after four
weeks at 25°C; on MEA black, velvety, partly immerse, raised at center with a
serrate 2.5 mm wide white edge band, radially sulcate, reverse dark gray; on
CMA mycelium white, inconspicuous, colonies low, plane, buff, pale gray at
center on both sides. Hyphae colorless, thin-walled, septate, smooth, some cells
moniliform. CONIDIOPHORES undifferentiated, single, short, colorless, smooth,
unicellular, reduced to conidiogenous cells. CONIDIOGENOUS CELLs intercalary
or terminal, colorless, cyathiform, (2.7—)3.8-5.4(-5.6) x (2.4-)2.8-3.6(-4.1)
um (n = 20), with a colorless multilayered cup-shaped collarette formed by
previously ruptured outer conidiogenous cell walls. Conip1A 1-celled, solitary,
smooth, thick-walled, translucent when immature, becoming dark brown
when mature, variably shaped, most ellipsoidal, ovoid, or pyriform with a germ
pore at the apical and basal ends, a few subspherical or even irregular, (9.5-)
10.5-13.5(-14.7) x (6-)6.3-7.3(-7.5) um (n = 30), some with truncate basal end
(2.2—)2.7-3.3(—4) um (n = 30) or apiculate, developing singly and successively
Bactrodesmiastrum domesticum sp. nov. (USA) ... 785
by percurrent extensions of the conidiogenous cell apices, secession schizolytic,
occasionally conidia developing directly on hyphae; mature conidia seceding
and released by rupture at the apex of the conidiogenous cell outer wall. Each
conidium leaves a layer of cup-like collarette in which a new conidium will
develop.
SPECIMEN EXAMINED: USA, MassaACHUSETTS, Hingham, wall ofa residence, 4 February
2013 (NHES L1702).
DISTRIBUTION: Czech Republic, Panama, USA.
SUBSTRATES/HOsTs: Dry wall, Scheelea zonensis (Corozo palm), decayed
wood of Ulmus glabra and an unidentified deciduous tree, and submerged
Ochroma pyramidale (balsa) wood (Piepenbring 2006; Shearer 1973).
Notes: Hohnel (1904) erected Conioscypha, typified by C. lignicola Hohn.
Shearer (1973), who described Conioscypha varia based on a specimen
collected from submerged balsa wood in the Patuxent River, MD, USA,
emended the generic concept of Conioscypha with detailed information
from her study on the conidiogenesis of C. lignicola. Réblova & Seifert (2004)
established Conioscyphascus, a new ascomycete genus in which both species,
Conioscyphascus varius (type species) and Conioscyphascus gracilis (Munk)
Réblova & Seifert, develop Conioscypha anamorphs. Their phylogenetic LSU
sequence analyses place C. japonica Udagawa & Toyaz., C. lignicola, and
Conioscyphascus varius in a monophyletic clade. Conioscypha (proposed in
1904 and with 17 species) has priority over Conioscyphascus (proposed in
2004 and with only two species) and is the generic name of choice for both the
asexual and sexual morphs under the “1Fungus :1Name’ principle. The C. varia
isolate collected indoors in Massachusetts did not develop the teleomorph.
Conioscypha varia is reported here as an indoor fungus for the first time. The
isolate collected indoors from drywall grew at a similar rateon CMA and MEA,
but produced no growth on DG18, indicating that C. varia is a hydrophilic,
not xerophilic, fungus. The holotype and four specimens of C. varia were all
collected during 1967-1969 from submerged balsa wood in Patuxent (Shearer
1973), a habitat indicating that it is an aquatic fungus. Its ecological characters
would make C. varia useful as an indicator fungus for long-term water damage
in indoor environments. After 40 years since this species was collected and
proposed as a new species, C. varia was once more collected in the USA, but
this time from an indoor environment and not from wood submerged in the
water.
It is possible that C. bambusicola Matsush. is conspecific with C. varia.
Conidia of these two species are similar in size. Conidia of C. bambusicola are
786 ... Li, Yang & Jalsrai
ovate and apiculate, while those of the C. varia holotype are variably shaped—
ovoid flammiform, naviculiform, subellipsoidal—with apiculate or rounded
apices and some conidia are ovoid, naviculiform, apiculate (Shearer 1973). The
shapes of the mature conidia from the isolate collected from the Massachusetts
indoor environment appeared similarly variable as those described by Shearer
(1973), while the immature conidia were mostly ovoid and apiculate, resembling
those described for C. bambusicola. Conidia from the isolate of C. varia studied
by Réblova & Seifert (2004) were ellipsoidal with a rounded apex. ‘The status,
delineation, and relationship of these two species should be further studied.
Acknowledgments
The authors are very grateful to Dr. Rafael F Castafieda Ruiz and Dr. Jian Ma and
for their critical review of the manuscript and to Drs. Theodore G. Andreadis and
James A. LaMondia for their pre-submission review. The authors are very appreciative
to Dr. Lorelei L. Norvell for her editorial review and Dr. Shaun Pennycook for his
nomenclature review. The authors are appreciative to Albert Baerren of Conestoga-
Rovers & Associates for providing the sample of Bactrodesmiastrum domesticum and
to Z-H Li for his assistance in obtaining literature otherwise unavailable to the authors.
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Holubova-Jechova V. 1972. Lignicolous hyphomycetes from Czechoslovakia - 2. Bactrodesmium.
Folia Geobotanica et Phytotaxonomica 7(4): 407-418. https://doi.org/10.1007/BF02854768
Holubova-Jechova V. 1984. Bactrodesmiastrum, a new genus of lignicolous hyphomycetes. Folia
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Mycological Research 103: 1409-1422. https://doi-org/10.1017/S0953756299008667
Manoharachary C, Agarwal DK, Sureshkumar G, Kunwar IK, Babu KS. 2006. Memnoniella
mohanramii sp. nov. and Zygosporium anupamvarmae sp. nov. from India. Indian
Phytopathology 59: 489-491.
Palm M, Stewart E. 1982. Two new combinations in Bactrodesmium. Mycotaxon 15: 319-325.
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Bactrodesmiastrum domesticum sp. nov. (USA) ... 787
Réblova M, Seifert KA. 2004. Conioscyphascus, a new ascomycetous genus for holomorphs with
Conioscypha anamorphs. Studies in Mycology 50: 95-108.
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Mycologia 65: 128-136. https://doi.org/10.2307/3757793
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Mycologia 63: 237-260. https://doi.org/10.2307/3757758
Yang J, Maharachchikumbura SSN, Bhat DJ, Hyde KD, McKenzie EHC, Jones EBG, Al-Sadi AM,
Lumyong S. 2016. Fuscosporellales, a new order of aquatic and terrestrial Hypocreomycetidae
(Sordariomycetes). Cryptogamie, Mycologie 37(4): 449-475.
https://doi.org/10.7872/crym/v37.iss4.2016.449
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 789-797
https://doi.org/10.5248/132.789
Russula brunneovinacea sp. nov., from northeastern China
XuU-MENG JIANG ©”, YANG-KUN LI?, JUN-FENG LIANG * & JIAN-RONG Wu ?
' Research Institute of Tropical Forestry, Chinese Academy of Forestry
Guangzhou, 510520, PB. R. China
? Southwest Forestry University, Kunming 650224, PR. China
* CORRESPONDENCE TO: jfliang2000@163.com
ABSTRACT—Russula brunneovinacea is described as a new species from northeastern China.
It is characterized by its deep brownish vinaceous to Etruscan red or Kaiser brown pileus with
a slightly depressed center, incurved margin with striate, Naples Yellow context that dries
with a greyish tinge, subglobose to broadly ellipsoid basidiospores with an amyloid plage
and isolated warts that are connected at base or in ridges warts but do not form a reticulum.
The new species is supported by ITS phylogenetic analyses and tentatively placed in Russula
subsect. Integroidinae based on morphological analyses.
KEY worps—macrofungi, Russulaceae, taxonomy
Introduction
Russula Pers. (Russulaceae, Russulales, Basidiomycota) was erected by
Persoon (1796) and with a worldwide distribution (Kirk et al. 2008). The
genus, which is ectomycorrhizal, includes many important edibles such as
R. griseocarnosa, popular in China and Southeast Asia (Wang et al. 2009, Li
et al. 2010, Zhao et al. 2015) as well as poisonous species such as R. emetica
(Schaeff.) Pers. and R. subnigricans Hongo (Miller & Buyck 2002, Yang &
Piepenbring 2004, Li et al. 2015).
Russula subsect. Integroidinae (Romagn.) Bon was validated by Bon (1986),
based on R. sect. Integroidinae Romagn. (Romagnesi 1958). Subsequently Bon
placed the subsection within R. sect. Lilaceae (Bon 1988). In this paper, we
follow the taxonomy of Bon (1986), who characterized species within this
subsection by a pileus with red, yellow, purple and complex color, a grayish
790 ... Jiang & al.
context with a rarely blackening tinge, pileocystidia that are absent or rarely
septate when present, and sometimes with primordial hyphae.
During our research on species diversity of Russula in the Changbai
Mountains of northeastern China, we identified two interesting Russula
collections as a novel taxon, which we tentatively place in R. subsect.
Integroidinae based on detailed morphological examination.
Materials & methods
Specimens, collected from Changbai Mountains, Jilin Province, during 2012, were
photographed in the field, with macroscopic descriptions of the fresh fruiting bodies
based on the photos and notes made in the field. After being dried at 50-55°C with an
electric drier, the collections were deposited at the Research Institute of Tropical Forestry,
Chinese Academy of Forestry Sciences, Guangzhou, China (RITF). Terminology for
descriptive terms follows Vellinga (1988). Color names and codes refer to Ridgway
(1912).
For microscopic observations, sections of specimens were cut by hand, mounted
in 5% KOH, and then stained with Congo red. We used Melzer’s reagent to determine
spore amyloidy and Cresyl Blue to test spores for a metachromatic reaction (Singer
1986). Sulfovanillin (SV) was used to test for reactions of cystidia. Sizes of basidia,
basidiospores, cheilocystidia, pleurocystidia and elements of the pileus covering were
measured using the MShot Digital Imaging System. The abbreviation [n/m/p] denotes
measurements made on ‘n basidiospores in ‘m’ basidiomata from ‘p’ collections.
Basidiospore dimensions follow the notation (a—)b-c(-d); the range ‘b-c’ includes 90%
of the measured values with the extreme values (‘a, ‘d’) given in parentheses. The spore
quotient Q indicates the length/width ratio of spores, and Qav denotes the average Q of
all basidiospores + sample standard deviation.
DNA was extracted from dried materials with an improved CTAB protocol (Zhou
& Liang 2011). Sequences of ITS region were amplified with the primers ITS5 and ITS4
(White et al. 1990). PCR products were purified using the Bioteke DNA Purification Kit.
Both strands were sequenced with an ABI 3730 DNA analyzer and an ABI BigDye 3.1
terminator cycle sequencing kit (Shanghai Sangon Biological Engineering Technology
& Services). These newly generated sequences were submitted to GenBank.
DNA sequences were edited and aligned by SeqMan (DNA STAR Package) and
MUSCLE (Edgar 2004) and manually checked and adjusted. Ambiguously aligned
positions were excluded from the matrix. Gaps in alignment were treated as missing
data. All unambiguous characters and character transformations were equally weighted.
A blast search of the query sequence in GenBank was performed to determine the
most closely related taxa for molecular identification. A dataset was generated from the
highest scored hits most relevant for identification, as well as from samples previously
used in the phylogenetic reconstruction of Russula (Miller & Buyck 2002). Russula
virescens (Schaeff.) Fr. and R. foetens Pers. were chosen as outgroup taxa.
Russula brunneovinacea sp. nov. (China) ... 791
Maximum likelihood (ML) and Bayesian analyses were used to generate a phylogeny.
The ML tree generation and bootstrap analyses were performed with the RAxML
BlackBox online server (Stamatakis et al. 2008). All parameters in RAXML analysis were
at default, statistical support values were obtained using nonparametric bootstrapping
with 1000 replicates, and trees obtained prior to convergence were discarded before the
consensus tree. A bootstrap proportion greater than 75% is considered significant. The
results were subsequently exported to Dendroscope for tree viewing and editing (Huson
et al. 2007).
The dataset was further analyzed with MrBayes 3.1 (Huelsenbeck & Ronquist 2005)
under a general-time reversible (GTR) model (nst = 6) and the parameter model was
selected by the Akaike Information Criterion (AIC) as the best-fit likelihood model with
Modeltest 3.7 (Posada & Buckley 2004). Bayesian analyses were run 6 chains for 1.0
million generations with sampling every 1000 generations. Trees were pooled together
and used to generate a 50% majority-rule consensus tree with branch lengths. Nodes
that were recovered more than 95% of the time were considered to have a significant
posterior probability.
Taxonomy
Russula brunneovinacea X.M. Jiang, Yang K. Li & J.F. Liang, & sp. nov. FIG.1
MycoBAnk MB 819154
Differs from Russula griseocarnosa by its deep brownish vinaceous to etruscan red or
kaiser brown pileus, barium yellow spore print, and smaller basidiospores with smaller
spines.
TypE—China, Jilin Province, Changbai Mountains, Lushuihe hunting ground, alt. 1000
m, 21 August 2012, Zhang Xin 268 (Holotype, RITF 2242; GenBank KY114148).
ETYMOLOGY—The Latin word “brunneovinacea” refers to the brownish vinaceous color
of the pileus.
BASIDIOMATA medium. PILEUS 5-6 cm in diameter, hemispherical when
young, expanding to applanate or concave with a slightly depressed center,
surface viscid when young and moist, glabrous when mature; Light Coral
Red (x115’b) to Coral Red (xm15’) when young, Dark Vinaceous (xxvit1”’) to
Mineral Red (xxvm11’’k) or Burnt Sienna (119k) at centre after maturity, margin
slightly incurved, sometimes undulate and dehiscent, obviously striate, Deep
Brownish Vinaceous (xxx1x5’”) to Etruscan Red (xxvit5”) or Kaiser Brown
(x1v9’k), often becoming Hay’s Russet (x1v7’k) to Russet (xv13’k) or Ocher
Red (xxv115’’i) to Prussian Red (xxv115’”"k) when dried. LAMELLAE White (LI)
to Cream Color (xv119’f), adnate, not forked, lamellulae absent, becoming
Chamois (xxx19”b) or Salmon Color (x1v9’d) with greyish tinge when dried.
STIPE 4.5-5 x 1-1.5 cm, central, cylindrical to subcylindrical, enlarged towards
the base, white, often tinged Hermosa Pink (11f) tinge when mature, Baryta
792 ... Jiang & al.
Fic. 1. Russula brunneovinacea (holotype, RITF2242). A. Basidiomata; B. Basidiospores;
C. Basidia; D. Pleurocystidia; E. Cheilocystidia; EF. Pileipellis
Yellow (1v21f) and tinged with Eosine Pink (11d) when dried, originally solid,
then stuffed with a bread-textured trama to irregular cavities, color unchanging
when injured. CONTEXT white (Li), Naples Yellow (xv119’d) with greyish
tinge when dried. Opor a little sweet. TasTE few acrid. SpoRE PRINT Barium
Yellow (xv123’d).
Basip1ospores [52/2/2] (6.1-)6.2-8.5(-8.8) x (5.1-)5.3-7.1(-7.8) um,
Q = (1.0—)1.03-1.31(-1.43) (Qav = 1.18 + 0.09), subglobose to broadly ellipsoid,
Russula brunneovinacea sp. nov. (China) ... 793
rarely globose or ellipsoid; ornamentation amyloid; warts bluntly conical to
subcylindrical, 0.7-0.9 um tall, isolated or connected at base or ridges, not
forming a reticulum; plage distinctly amyloid; hyaline in 5% KOH. Basip1a
32-46(-52) x 10-13.5 um, narrowly clavate to clavate, inflated towards upper
half, 4-spored, rarely 2-spored, hyaline in KOH, sterigmata about 2.5-4.5 um
long. LAMELLAR TRAMA mainly composed of nested sphaerocytes (28-41 x
26-33 wm) surrounded by connective hyphae. PLEUROCYSTIDIA 54-89 x
9-13 um, abundant, narrowly clavate to clavate, apex often with papillate
appendage, with abundant granular contents in the upper part, red to slightly
purplish red and weakly grey in SV. CHEILOcysTIDIA 43-82 x 7-13 um, clavate
with rounded or indistinctly mucronate apex, few with granular contents.
PILEIPELLIS composed of hyaline hyphae (2-5 um diam.), often branched and
interwoven, septate; terminal cells 16-37 x 2.2-4.1 um, cylindrical, with obtuse
apex, sometimes attenuate, and some with zebroid encrustations. CLAMP
CONNECTIONS absent.
HABITAT & DISTRIBUTION — Solitary or dispersed in conifer and conifer-
broadleaf forests. So far found only from the type locality.
Additional specimen EXAMINED — CHINA: JILIN PROVINCE, Changbai Mountains, alt.
1000 m, 19 August 2012, Zhang Xin 226 (RITF 2206; GenBank KY114147).
Phylogenetic analyses
A dataset of 32 ITS sequences with 666 nucleotide sites (including 30
available sequences of Russula from GenBank) was analyzed. Both the RAxML
and Bayesian analyses resulted in similar topologies and only the best maximum
likelihood tree with bootstrap supports is shown (Fic. 2). Bayesian posterior
probability is also displayed with the bootstrap values along the branches.
Bayesian and RAxML phylogenetic analyses clustered two collections
together and obviously separate from the related sequences available
on GenBank. Phylogenetic analyses also grouped the new species with
R. griseocarnosa X.H. Wang et al., R. occidentalis Singer, R. vinosa Lindblad,
and other species with weak statistical support.
Discussion
Russula brunneovinacea is well distinguished by its Deep Brownish Vinaceous
to Etruscan Red or Kaiser Brown pileus with a slightly depressed center, its
incurved striate margin, its white or cream lamellae that become Chamois with
a greyish tinge when dried, its Naples Yellow context with greyish tinge when
dried, its somewhat acrid taste, and its subglobose to broadly ellipsoid spores
with small isolated or connected at base or ridges warts.
794 ... Jiang & al.
Bayesian and RAxML phylogenetic analyses support R. brunneovinacea
as closely related to the Chinese specimens previously misdetermined as
“R. vinosa” and subsequently described as a new species, R. griseocarnosa
(Wang et al. 2009). Russula griseocarnosa differs from R. brunneovinacea by
its dark red sticky pileus, white context turning greyish during maturity, white
spore print, stipe tapered at base, larger basidiospores (8-10 x 6.5-8 um)
with longer (<2.5 um) spines, and pleurocystidia (100-150 x 13-20 um).
Geographically, R. griseocarnosa invariably grows in tropical and subtropical
China while R. brunneovinacea grows in northeastern China with a temperate
monsoon climate.
Russula occidentalis is also close to R. brunneovinacea. A BLAST query
showed that the two species have a 92% maximum identification with 91%
coverage. However, R. occidentalis has a variably colored pileus, lamellae with
forking edges that bruise grey to black, a white to creamy white stipe that stains
dingy pink where handled or damaged and becomes dark grey to brownish-grey
or black, and bigger spores (7.9-11.2 x 6.7-8.9 um), basidia (52-62 x 14-17.7
um), and pleurocystidia (70-125 x 9-13 um) (Romagnesi 1985, Roberts 2007).
Russula vinosa has a 90% maximum identification with R. brunneovinacea;
it differs in its vinaceous red, dark wine-red or somewhat brownish pileus,
lamellae becoming grey to black when cut or bruised, pale ochre spore print,
mild taste and distinctive odor (Romagnesi 1967, Bon 1988).
In the field, R. brunneovinacea is easily confused with the United States
species R. rubescens Beardslee (Romagnesi 1985). Russula rubescens also has
red color on the pileus, but differs by its paler margin fading with age, mild taste,
forked lamellae, white stipe becoming cinereous, and often blackening with
age or upon drying, and quickly becoming red and then black when wounded,
and basidiospores with an indistinctly amyloid plage. Russula xerampelina
(Schaeff.) Fr. and R. olivacea (Schaeff.) Pers. (Romagnesi 1985, Roberts
2007) also resemble R. brunneovinacea in appearance. Russula xerampelina is
distinguished by its ochraceous-olive young pileus, non-striate margin, warm
yellow to light orange-yellow lamellae often staining brown where damaged,
cream to light yellow stipe, crab or shrimp odor, light orange yellow spore print,
and basidiospores with longer (<1.2 um long) warts, while R. olivacea differs
in its bigger (10-20 cm) basidiomata, drab pileus with olive green and purple
hues, non-striate margin, egg-yolk yellow lamellae unchanging when damaged,
deep ochre-yellow spore print, and basidiospores with incompletely amyloid
plage (Romagnesi 1985, Roberts 2007).
Russula brunneovinacea sp. nov. (China) ... 795
R.eccentrica KF306039
74 R.katarinae KP966377
R.favrei EF530944
R.pascua AY061705
75 R.xerampelina FJ845433
R.sphagnophila AY 061719
= ‘R.nitida KT934001
84/1.00
R.melliolens AY061690
‘R.subsulphurea KF810135
R.puellula JF908706
67/0.99 ‘R.lilacea AY061731
R.azurea AY (061660
R.rosea AY061715
76/0.99 R.rubra AY061717
100 R.amethystina AY 061653
R.firmula KJ867372
R.californiensis AY 245542
R.decolorans FJ845432
100/1.00) R.brunneovinacea KY114148
66/0.99 R. brunneovinacea KY114147
‘R. griseocarnosa EF627042
R.claroflava AY 061665
R.vinosa AY061724
77/100 “R.occidentalis AY 534206
R.velenovskyi AY061721
R.integra AY 061683
96/1.00 R.integriformis AY 061684
R.lepida AY 061686
‘R.aurantiaca AY 061658
R.paludosa AY061703
R foetens AY061677
R.virescens AY061727
60
64
89/1.00
69
pe ON en nstitiionieite
Fic. 2. The best RAxML likelihood tree (-In L 3803.544073) based on the Russula ITS dataset.
Support values in bold type are RAxML likelihood bootstrap (250%). Values in normal type are
Bayesian posterior probabilities (20.95).
796 ... Jiang & al.
Acknowledgments
We are grateful to Dr. Yanchun Li and Dr. Baokai Cui for their critical reviews of
the manuscript. This work was supported by the National Natural Science Foundation
of China (No.31570544), Natural Science Foundation of Guangdong Province
(2014A030313727), Forestry Science and technology innovation Fund of Guangdong
(2014KJCX019-01) and the National Nonprofit Institute Research Fund of CAF
(CAFYBB2014MA003).
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 799-811
https://doi.org/10.5248/132.799
Amanita pallidorosea in Pakistan and
its ectomycorrhizal association with Quercus oblongata
MUNAZZA KIRAN’ , JUNAID KHAN’, AROoj NASEER’,
HASSAN SHER’ & ABDUL NASIR KHALID'
University of the Punjab, Department of Botany,
Quaid-e-Azam Campus-54590, Lahore, Pakistan
*University of Swat, Center for Plant Sciences and Biodiversity,
Saidu Sharif-19200, Swat, Pakistan
* CORRESPONDENCE TO: munazzakiran@gmail.com
ABSTRACT—The occurrence of Amanita pallidorosea in Pakistan and its ectomycorrhizal
association are reported for the first time based on morphological characters and nucleotide
sequences of the internal transcribed spacer region generated from basidiomata and from
ectomycorrhizal roots of Quercus oblongata [= Q. incana Roxb., nom. illeg.].
Key worps—Amanitaceae, ITS, lethal, phylogeny, Swat
Introduction
Amanita Pers. (Agaricales, Basidiomycota) is a large genus comprising
about 500 described species (Bas 2000, Tulloss 2005, Yang 2005, Kirk et al.
2008). Many Amanita species have been shown to be ectomycorrhizal, thus
playing vital roles in forest ecology (Yang 1997). New Amanita species have
been regularly reported from all over the world, including East Asia, Central
and South America, South Africa, and Australia (Zhang et al. 2015). However,
knowledge about the genus is rather limited for Pakistan, where only eighteen
species have been reported (Ahmad et al. 1997, Tulloss et al. 2001). The goal of
this study was to collect Amanita basidiomata and ectomycorrhizae from the
Quercus-dominated forests of Swat district, Khyber Pakhtunkhwa province,
Pakistan. Mycological expeditions took place during the monsoon season
(July to September) of 2015. Nuclear ribosomal DNA sequence analyses of
800 ... Kiran & al.
the ITS region from basidiomata and ectomycorrhizal roots were combined
with detailed morphological examinations to confirm Pakistani collections as
A. pallidorosea, a new record for Pakistan. We describe the ectomycorrhiza of
this species here for the first time.
Materials & methods
Sampling site
Different sites dominated by oak forests were selected for sampling within district
Swat (34°34’-35°55’N 72°08’-72°50’E). The Swat valley, well known for its biodiversity,
is in Pakistan's Khyber Pakhtunkhwa (KP) province (Shinwariet al. 2003) where altitudes
range from 970 m to 2300 m (Barth 1956), with Falak Sar (the highest peak) towering
5918 m above sea level. The district’s climate is dry temperate to moist temperate. Most
of the region gets monsoon rains in summer except in the north, which receives very
little or no summer monsoon rains (Ali & Qaisar 1986, Ahmad & Sirajuddin 1996). Swat
vegetation types range from sub-tropical chir pine (Pinus roxburghii Sarg.) forest in the
south to alpine pastures and meadows or cold desert in the north (Sher 2013).
Collection and morphological characterization of the basidiomata
Basidiomata were collected, tagged and photographed in the field using a Nikon
D70S digital camera. Fresh morphological characters were recorded in the field and
colors were designated using Munsell (1975) after which the specimens were dried with
the help ofa fan heater and sealed in zip-lock bags for long-term preservation. The dried
material was vouchered and deposited in the Lahore Herbarium, Department of Botany,
University of the Punjab, Pakistan (LAH).
Microscopic characteristics were observed in 5% KOH solution. The abbreviation
‘n/b/p’ indicates ‘n’ basidiospores measured from ‘b’ basidiomata from ‘p’ collections.
Basidiospore dimensions are given as length x width (1 x w), with extreme values given
in parentheses, and Q = I/w ratio of individual spores; avQ = average Q of all spores +
standard deviation (Zhang et al. 2010).
Isolation and morphological characterization of ectomycorrhizal roots
Twenty 15 cm’ soil cores near the soil surface were retrieved from each sampling site
close enough to Quercus tree trunks to ensure that they contained fine Quercus roots.
Each soil core was later soaked in water in the laboratory for 1-2 hrs and then passed
over a 2 mm sieve under running water to separate the roots from soil. Ectomycorrhizal
roots were carefully sorted into morphotypes using a Meiji Techno EMZ-5TR
stereomicroscope. The fresh ectomycorrhizal roots were cleaned with a fine camel hair
brush, photographed under the stereomicroscope, and examined anatomically under a
compound microscope. The morphotypes were kept in McCartney bottles in distilled
water with replicates of each reserved in 2% CTAB buffer at 8°C for molecular analysis.
Amanita pallidorosea in Pakistan ... 801
DNA extraction, PCR and sequencing
DNA was extracted (from 3-4 gills or 0.5-1 mg of context tissue per basidioma
or 2-4 ectomycorrhizal root tips per morphotype) using a modified CTAB method
(Bruns 1995). The extracted genomic DNA was evaluated using gel electrophoresis
(1% agarose gel in Gel documentation system Uvipro Platinum (IM-2000 WL/
LC/26M MANZ) with default settings. After confirmation, genomic DNA was
suspended in nuclease free water and stored at 20°C. The target rDNA internal
transcribed spacer region (ITS1 + 5.8S + ITS2) was amplified by optimized polymerase
chain reaction (PCR) (Gardes & Bruns 1993) using the ITS1F fungal specific forward
primer (5’-CTTGGTCATTTAGAGGAAGTAA-3'; Gardes & Bruns 1993), ITS4 reverse
primer (5’-TCCTCCGCTTATTGATATGC-3'; White et al. 1990), and Taq polymerase as
catalyst in 20 uL PCR tubes using Gene Amp PCR system 9700. The thermo-cycling
profile had an initial and final denaturation step at 94°C for 1 minute followed by
30 cycles at 53°C for 1 minute for annealing, and a 1-minute initial extension and
8-minute final extension at 72°C. PCR products were visualized by using 1% agarose
gel. Amplified products were sequenced in both directions with the ITS1r and ITS4
amplification primers by BGI Tech Solutions (HK Co. Ltd.). To identify the plant
host species, the host plant DNA was extracted from root tips and amplified using
the plant specific primer 28xky (5’-GGCGGTAAATTCCGTCC-3’) and universal primer
28c (5’-GCTATCCTGAGGGAAACTT-3’) (Cullings 1992). Successful amplicons obtained
from ectomycorrhizal roots were purified using Exonuclease I and Shrimp Alkaline
Phosphatase enzymes (Werle et al. 1994). The purified products were sequenced by
the University of Florida's Interdisciplinary Center for Biotechnology Research (http:
//www. biotech. ufl.edu/). Sequence chromatograms were trimmed, edited, and
assembled using Sequencher 4.1 (Gene Codes, Ann Arbor, MI).
Sequence alignment and phylogenetic analysis
Consensus sequences were generated from the obtained sequences with the forward
and reverse algorithms in BioEdit (Hall 1999) and then BLAST searched at NCBI (http://
www.ncbi.nlm.nih.gov/). Closely matched sequences from GenBank were downloaded
for phylogenetic analysis based on BLAST analysis and on sequences generated by
previous phylogenetic studies of Amanita pallidorosea (Zhang et al. 2010, Kim et al.
2013, Li et al. 2015). Sequences were manually edited and assembled using BioEdit (Hall
1999) and ClustalW (Thompson et al. 1997). All ITS sequences were trimmed with the
conserved motifs 5’-(...GAT)CATTA— and -GACCT(CAAA...)-3° to facilitate alignment,
with the the alignment portion between them included in the analysis (Dentinger et al.
2011). These sequences were aligned with other GenBank sequences using the MUSCLE
(Edgar 2004) alignment tool available online (www.ebi.ac.uklTools/msalmuscle). In
aligned dataset, all characters were equally weighted, and gap positions were treated as
missed data. A phylogenetic tree was constructed with the maximum likelihood (ML)
analysis using Jukes-Cantor model in MEGA6 software to test the phylogeny (Tamura et
al. 2011). All obtained sequences have been submitted to and accessioned by GenBank.
802 ... Kiran & al.
Results
Amanita pallidorosea P. Zhang & Zhu L. Yang,
Fungal Diversity 42: 125 (2010) Fics 1, 2
BASIDIOMATA medium sized. PILEUs 40-60 mm diam., convex to plano-
convex, slightly umbonate, the disc pale pink (5yR9/4) to pale rose (10R9/6) and
becoming white with maturity or under dry conditions; surface dry, smooth,
volval remnants sometimes present, dull to shiny, margins smooth, incurved
when young becoming straight and splitting with age or in dry conditions;
context dry, whitish or slightly paler than cap, 1-3 mm, texture soft, not
changing upon cutting or bruising. OpoR not distinctive; taste not recorded.
LAMELLAE 4.5 mm broad, free, crowded, whitish at first then turning creamy
to yellowish cream (10YR9/8) at maturity, edge entire. LAMELLULAE attenuate,
frequent, length variable. Stipe 80-120 x 6-12 mm, central, cylindrical to sub-
cylindrical to tapering upwards, white to creamy white above the annulus and
creamy white with a yellowish (10yYR9/8) to yellowish pink (10R7/10) tinge
below, upper quarter hollow, below solid; surface smooth to finely fibrillose
squamulose; context white. ANNULUS present, membranous, thin, permanent,
yellow to white. VoLva saccate, white, membranous, permanent, in some
specimens breaking to leave fragments on the stipe, giving the stipe a banded
appearance.
Basipiospores [n/b/p = 80/4/3] 7-8.5(-9) x (6.5-)7-8.5(-9) um, Q =
1-1.08, avQ = 1.01, globose to subglobose, apiculus prominent (1.5 um long),
guttulate, amyloid in Melzer’s reagent. BAsip1A 31-40 x 8-10.5 um, clavate,
tetra-sterigmate, basal clamps absent. PILEIPELLIs hyphae 3.5-5(-7.5) um
diam., filamentous, terminal inflated cells clavate 15 um diam., hyaline; when
present, volval remnant cells abundant, inflated (18.5 x 18 um). STIPITIPELLIS
filamentous, hyaline, 4.5-12 um diam. VOLVAL CELLS subglobose to oval (66
x 78.5 um), hyphae 5-7 um diam. CLAMP CONNECTIONS absent in all tissues.
SPECIMENS EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA PROVINCE, Swat
District, Lower Shawar, 2100 m asl, on ground in a forest dominated by Quercus
oblongata, 5 September 2015, Zia-ullah Sh0O3 (LAH35204; GenBank KY626178);
Malam Jabba, Kishawra, 2200 m asl, on soil under oak, 16 August 2015, Junaid Khan
MJ1554 (LAH35205; GenBank KY621476); Toa, 2800 m asl, on ground under Quercus
oblongata, 15 July 2015, Arooj Naseer T4 (LAH35206; GenBank KY616968).
COMMENTS: Macroscopically, Amanita subpallidorosea Hai J. Li is strikingly
similar to A. pallidorosea because of its pale rose pileus with a conspicuous
umbo. However, microscopically, A. subpallidorosea can be distinguished
from A. pallidorosea by its larger basidia and basidiospores (Li et al. 2015,
Yang 2015). Pure white specimens of A. pallidorosea, also common (Zhang et
Amanita pallidorosea in Pakistan ... 803
: yy lad ao /
Fic. 1. Amanita pallidorosea basidiomata. A. LAH35204; B. LAH35206; C-F LAH35205.
Scale bars: A = 1.9 cm; B =1.6 cm; C = 0.9 cm; D = 1.1 cm; E= 1.7 cm; F= 1.5 cm.
804 ... Kiran & al.
Fic. 2. Amanita pallidorosea (LAH35204). A. basidia and subhymenium; B. volval remnants on
pileus; C. basidiospores; D. volva; E. pileipellis; F. stipitipellis. Scale bars: A = 7.5m; B = 30 um;
C= 6.5 um; D = 25 um; E, F = 20 um. Drawings by M. Kiran.
al. 2010, Yang 2015), can be easily confused with A. virosa Bertill. (Malysheva
et al. 2014). Amanita virosa, however, can be distinguished with its distinctly
recurved squamules on the stipe and larger basidiospores (Neville & Poumarat
2004, Yang 2015).
Amanita pallidorosea in Pakistan ... 805
Fic. 3. Amanita pallidorosea/Quercus oblongata mycorrhiza (LAH-EM64). A, B. ectomycorrhizal
roots; C. inner mantle layer cells; D. outer mantle layer cells. Scale bars: A = 0.6 mm; B = 1.2 mm;
C=7.5 um; D = 10 um. Drawings by M. Kiran.
Morphological characterization of ectomycorrhizae FIG. 3
ECTOMYCORRHIZAL SYSTEM dichotomous, 6-18 mm long, axis <1 mm
diam., unramified, ends straight, cylindrical, surface smooth, dark brown
806 ... Kiran & al.
(10R1/4). OUTER MANTLE hyphal cells 4.39-5.85 um diam., loosely arranged.
INNER MANTLE hyphal cells 4.17-4.80 um compactly arranged. Cells in both
layers are mantle type A (Agerer 1987-2002), frequently septate, branched,
thin-walled, yellowish brown, end cells not sharply pointed. EMANATING
HYPHAE rarely present. RHIZOMORPHS not observed.
HABITAT & DISTRIBUTION: In Pakistan, solitary to scattered in small groups
on ground under oak. Originally described from China (Zhang et al. 2010);
also known from Japan, Korea, and eastern Russia (Kim et al. 2013, Malysheva
et al. 2014, Yang 2015, Cai et al. 2016).
SPECIMENS EXAMINED: PAKISTAN, KHYBER PAKHTUNKHWA PROVINCE, Swat
District, Toa, 2800 m asl, associated with roots of Quercus oblongata, 15 July 2015,
Arooj Naseer ANT58 (LAH-EM65; GenBank KY616969); 15 July 2015, Arooj Naseer
ANT81 (LAH-EM66; GenBank KY616970); 15 July 2015, Arooj Naseer ANT450044
(LAH-EM67; GenBank KY616971); Shawar, 2100 m asl, on roots of Quercus oblongata,
14 August 2015, Arooj Naseer AN26 (LAH-EM64; GenBank KY616967).
Molecular phylogenetic characterization Fic. 4
The host plant sequence (GenBank KY829452) obtained from
ectomycorrhizal roots of Amanita pallidorosea (GenBank KY616969) showed
99% identity with Quercus oblongata D. Don [= Quercus incana Roxb., nom.
illeg.] (GenBank AY456170) using BLAST search tool at NCBI. Phylogenetic
analysis was performed on a 58-sequence ITS dataset of which 57 sequences
represented Amanita species and one represented the outgroup Limacella
glioderma (Fr.) Maire [= L. delicata (Fr.) Konrad & Maubl.]. The post-alignment
dataset included 891 sites, of which 339 were conserved, 489 variable, and 345
parsimony informative. Reliability of the consensus tree was calculated by 1000
bootstrap replications. The phylogenetic tree contained four well supported
clades corresponding to four Amanita sections: A. sect. Phalloideae, A. sect.
Validae, A. sect. Lepidella, and A. sect. Amanita (Zhang et al. 2010, Cai et al.
2016). Sequences obtained from the Pakistani basidiomata and morphotypes
clustered with A. pallidorosea in A. sect. Phalloideae. This lineage is consistent
with the morphological identification.
Discussion
Amanita pallidorosea is one of the lethal amanitas in A. sect. Phalloideae
(Zhang et al. 2010, Malysheva et al. 2014, Yang 2015). First described by
Zhang et al. (2010) from China, the species is characterized by typically pale
rosy (sometimes white) umbonate pileus and small globose to subglobose
basidiospores. This taxon has been reported from other Asian countries
Amanita pallidorosea in Pakistan ... 807
i Amanita pallidorosea KY616967
@ Amanita pallidorosea KY616968
i Amanita pallidorosea KY616971
Amanita pallidorosea KJ466382
Amanita pallidorosea FJ176736
66] Hl Amanita pallidorosea KY616970
Mi Amanita pallidorosea KY616969
Amanita pallidorosea KF245917
Amanita pallidorosea KF245915
oe Amanita pallidorosea KT894838
@ Amanita pallidorosea KY626178
do! @ Amanita pallidorosea KY621476
Amanita pallidorosea KF245916
Amanita pallidorosea KJ466389
Amanita pallidorosea KT894837
98] + Amanita subjunquillea var alba FJ375332
Amanita pallidorosea KJ466383
Amanita pallidorosea JX998035
Amanita pallidorosea JX998037
Amanita pallidorosea KJ466385
Amanita bisporigera KU311692
gg ' Amanita bisporigera KRQ19764
Amanita rimosa KU311696 : ;
Toot Amanita rimosa KU311695 | “tion Phalloideae
7-— Amanita virosa KF937304
63 Amanita virosa ABO15676
84, Amanita subpallidorosea KP691682
691 |! Amanita subpallidorosea KP691678
gg | Amanita subpallidorosea KP691680
gi! Amanita subpallidorosea KP691677
Amanita ocreata KJ466381
gq ‘Amanita ocreata GQ486374
94, Amanita phalloides GQ221843
Amanita phalloides GQ221841
ge |r Amanita subjunquillea KR996715
e9' Amanita subjunquillea KT894848
99 - Amanita suballiacea KP221316
Amanita suballiacea KP221303
100; Amanita fuligineoides FJ176721
90 Amanita fuligineoides FJ176720
Amanita reidii AY325824
Amanita exitialis KT213706
80 Amanita exitialis KT213705
Amanita ocreata AY918962
Amanita areolata AB167727
gg ‘Amanita zangii KJ466433
Amanita spissa KF245910
Amanita rubescens ABO15682 _| Section Validae
9g + Amanita rubescens EU819464
Amanita cylindrispora AY325839
57 Amanita oberwinklerana KT894853 —_| Section Lepidella
100 ' Amanita oberwinklerana KT894852
100 , Amanita subglobosa KU139498
Amanita subglobosa KU139497
Amanita subfrostiana JN9431 72 Section Amanita
99 Amanita rubrowlvata FJ375329
75 Amanita subfrostiana KF651010
100
91
Limacella glioderma FJ478086 “Joutgroup
0.05
Fic. 4. Molecular phylogenetic analysis of Amanita spp. based on ITS sequences. The tree with
the highest log likelihood (—5886.3380) is shown. The analysis involved a total of 58 nucleotide
sequences and 891 positions in the final dataset. New sequences generated from Pakistani
basidiomata are marked with @ and from ectomycorrhizal root tips with Ml.
808 ... Kiran & al.
including Japan (Yang 2015, Cai et al. 2016), Korea (Kim et al. 2013, Cho et
al. 2015), and Russia (Malysheva et al. 2014). Macro- and microscopically, the
three A. pallidorosea basidiomata (LAH35204-35206) collected in Pakistan
agree with the descriptions by Zhang et al. (2010) and Malysheva et al. (2014).
However, the smooth stipe surface in some specimens and the hollow stipe
apex contrast with the original description. The yellowish annulus in some
collections, although differing from the original collection of Zhang et al.
(2010), conforms to the findings of Malysheva et al. (2014) and Yang (2015).
Amanita pallidorosea has not been previously recorded from Pakistan (Ahmad
et al. 1997), and our basidiomata and ectomycorrhizal roots represent the first
report of the species from the country. Ectomycorrhizal association of Amanita
muscaria (L.) Lam., A. citrina Pers., and A. strobiliformis (Vittad.) Bertill. have
been described with various hosts with different ramification patterns (Ingleby
et al. 1990, Mleczko 2004, Raid] & Verma 2006)
Phylogenetically, the sequences obtained from ectomycorrhizal roots
and basidiomata clustered with A. pallidorosea sequences retrieved from
GenBank in clade sect. Phalloideae, which contained A. areolata T. Oda et
al., A. bisporigera G.F. Atk., A. exitialis Zhu L. Yang & T.H. Li, A. fuligineoides
P. Zhang & Zhu L. Yang, A. ocreata Peck, A. phalloides (Fr.) Link, A. reidii
Eicker & Greuning, A. rimosa P. Zhang & Zhu L. Yang, A. suballiacea (Murrill)
Murrill, A. subjunquillea S. Imai, A. subjunquillea var. alba Zhu L. Yang,
A. subpallidorosea, A. virosa, and A. zangii Zhu L. Yang et al. (Cai et al. 2014,
2016). Large white basidiomata morphologically similar to A. pallidorosea are
also present in A. bisporigera, A. exitialis, A. oberwinkleriana Zhu L. Yang &
Yoshim. Doi, A. ocreata, A. subjunquillea var. alba, and A. virosa (Imazeki &
Hongo 1987, Yang & Li 2001, Malysheva et al. 2014) (Fic. 4).
Amanita pallidorosea has been previously reported from forests dominated
by Quercus mongolica Fisch. ex Ledeb., and members of family Fagaceae and
Pinaceae (Zhang et al. 2010, Malysheva et al. 2014, Cai et al. 2016), but no
studies have been published confirming its ectomycorrhizal association. This
appears to be the first report of ectomycorrhizal association of A. pallidorosea
with Quercus oblongata supported by morphological and molecular data.
Acknowledgements
The authors are grateful to Prof. Zhu Liang Yang (Kunming Institute of Botany,
Chinese Academy of Sciences, China) for his valuable comments and suggestions to
improve the manuscript. Thanks are due to Dr. Sana Jabeen (University of Education,
Faisalabad Campus, Pakistan), Dr. Andrew W. Wilson (Denver Botanic Gardens, USA),
Amanita pallidorosea in Pakistan ... 809
and Dr. Yan-Chun Li (Kunming Institute of Botany, Chinese Academy of Sciences,
China) for providing helpful reviews of this manuscript. We very much appreciate the
field assistance of Dr. Abdul Rehman Khan Niazi (University of the Punjab, Lahore,
Pakistan). Sincere thanks are also due to Dr. Matthew E. Smith (University of Florida,
USA) for providing molecular research facilities to one of the authors. Thanks are also
due to Dr. Shaun Pennycook for his precious time in editing the nomenclature.
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 813-817
https://doi.org/10.5248/132.813
Ellisembia pseudokaradkensis sp. nov.
from Hainan, China
MIN QIAO, XING Du, ZHAO-Hut!I BIAN, JIE PENG & ZE-FEN YU*
Laboratory for Conservation and Utilization of Bio-resources,
Key Laboratory for Microbial Resources of the Ministry of Education,
Yunnan University, Kunming, Yunnan, 650091, P. R. China
* CORRESPONDENCE TO: Zfyuqm@ynu.edu.cn
ABSTRACT—Ellisembia pseudokaradkensis, isolated from decaying dicotyledonous
leaves submerged in a stream, is described and illustrated as a new species. The fungus is
characterized by obclavate conidia with long flagelliform apical appendages. The new taxon
is compared with other Ellisembia species having flagelliform appendages at the conidial tip.
Key worps— aquatic hyphomycetes, fungal diversity, morphology
Introduction
Ellisembia was introduced by Subramanian to accommodate Sporidesmium-
like species, with Sporidesmium coronatum Fuckel [= E. coronata (Fuckel)
Subram.] as type species (Subramanian 1992). The genus is characterized
by determinate or irregularly percurrently extending conidiogenous cells
that produce distoseptate conidia, while Sporidesmium is restricted to those
species that have conidiogenous cells without or with irregular proliferation
and euseptate conidia (Subramanian 1992). Despite the segregation of a new
genus from Sporidesmium in 1992, researchers continued to place some species
with distoseptate conidia into Sporidesmium (Shi & Zhang 2007, Ma & Zhang
2007a,b); these were subsequently transferred into Ellisembia (Santa-Izabel
et al. 2013). Wu & Zhuang (2005), who merged Imicles Shoemaker & Hambl.
into Ellisembia, expanded the generic concept to include species that produce
typically lageniform, ovoid, or doliiform, percurrently extending conidiogenous
cells. Currently 60 species are included in Ellisembia (Index Fungorum 2017).
814 ... Qiao & al.
Several fungal species and genera occurring on dead stems or decaying
branches of broadleaf trees have been described recently in China (Xia et al.
2015, 2016; Ren et al. 2012; Ma et al. 2007a,b, 2010). During a study of aquatic
hyphomycetes in Diaoluo Mountain Preserve, Hainan Province, China, we
encountered an unknown fungus sharing some features of Ellisembia that we
propose here as a new species.
Materials & methods
The culture was isolated from decaying leaves collected on a broadleaf tree in
Diaoluo Mountain Preserve, Hainan province, in southern China, by Z.E Yu on
April 2015. The decaying leaves were cut into several 2-4 x 2-4 cm sized fragments,
which were flattened out on the surface of CMA (20 g cornmeal, 18 g agar, 40 mg
streptomycin, 30 mg ampicillin, 1000 ml distilled water) for ten days; single conidia
were isolated using a sterilized toothpick while viewing with a CX31 microscope,
and cultivated on CMA in Petri plates. Morphological observations were made from
CMA after incubation at 28°C for one week; pure cultures and a permanent slide
were deposited in the Herbarium of the Laboratory for Conservation and Utilization
of Bio-resources, Yunnan University, Kunming, Yunnan, P.R. China (YMF; formerly
Key Laboratory of Industrial Microbiology and Fermentation Technology of
Yunnan).
Taxonomy
Ellisembia pseudokaradkensis M. Qiao & Z.F. Yu, sp. nov. PLATE 1
MycoBank MB 820988
Differs from Ellisembia karadkensis by its smaller, unbranched conidia with fewer septa.
Type: China, Hainan Province: Diaoluo Mountain Preserve, 18°43’N 109°43’E, elev.
1020 m, on submerged decaying leaves of an unidentified dicotyledonous plant in a
stream, July 2015, J. Peng (Holotype, YMF 1.04266).
EryMo oey: Latin, pseudokaradkensis, referring to the similarity to E. karadkensis in
the shape of conidia.
CoLonigs 2 cm diam. on CMA after 10 days, Mycelium partly superficial and
partly immersed in the substratum. CONIDIOPHORES erect, macronematous,
mononematous, straight or flexuous, dark to blackish brown, arising from
creeping hyphae, 0-1-septate, 7-18 x 4-5 um, base often specialized to T- or
L-shape. CONIDIOGENOUS CELLS monoblastic, integrated, terminal, cylindrical,
pale or dark brown, 14-15 x 4.5 um. Conidial secession schizolytic. CONIDIA
solitary, obclavate, mostly curved, tapering towards narrow whip-like apical
tip, (81.5-)90.5-138(-149) um long, 5.5-8 um diam., truncate, 2.5-4 um diam.
at the base, 8-13-distoseptate, with 2-3 apical cells subhyaline or hyaline with
Ellisembia pseudokaradkensis sp. nov. (China) ... 815
PiaTE 1. Ellisembia pseudokaradkensis (holotype, YMF 1.04266). A. Distoseptate conidium.
B. Conidia with flagelliform apical appendages. C. Conidiophores and conidiogenous cells.
Scale bars: A, C = 10 um; B = 20 um.
816 ... Qiao & al.
thin septa, other cells pale olivaceous to brown but dark at the septa, apical
appendages (33-)57-148 um long, 0.5-1 um diam.
Discussion
Ellisembia pseudokaradkensis is characterized by flagelliform apical
appendages and long, narrow conidia. Five other Ellisembia species have
flagelliform appendages at the tip of conidia: E. podocarpi Jian Ma &
X.G. Zhang, E. photiniae Jian Ma & X.G. Zhang, E. flagelliformis (Matsush.)
W.P. Wu, E. magnibrachypus (Matsush.) Rajeshk. & S.K. Singh, and
E. karadkensis Rajeshk. & S.K. Singh. Ellisembia E. karadkensis is
distinguished from E. pseudokaradkensis by its branched conidia and
E. magnibrachypus by its branched appendages, and E. pseudokaradkensis
is distinguished from all five species by its distinctive combination of the
number of conidial septa, conidial size, and length of apical appendage
(TABLE 1).
TABLE 1. Comparison of Ellisembia species with appendages at the tip of conidia.
CONIDIA
SPECIES ee ,._—0. 52 OE REFERENCE
NUMBER OF LENGTH X APICAL APPENDAGE
SEPTA WIDTH (tm) LENGTH (um)
E. flagelliformis 11-14 60-90 x 40 Wu & Zhuang
9-10 (2005)
E. karadkensis 10-35 150-345 x 100 * Rajeshkumar et al.
9.5-12.5 (2012)
E. magnibrachypus 9-10 48-62 x 65 Matsushima (1975)
12-14
E. photiniae 10-16 92-170 x 43-90 Ma et al. (2010)
13-16
E. podocarpi 13-19 110-170 x 80 Ma et al. (2010)
7.5-10
E. pseudokaradkensis 8-16 90.5-138 x (33-)57-148 Thi
6.0-7.5 v8 Paper
* Appendage length estimated from illustration
Acknowledgements
This work was jointly financed by National Natural Science Foundation Program
of PR China (31360130, 31770026). We are very grateful to Prof. X.G. Zhang and
Dr. R.E Castafieda-Ruiz for critically reviewing the manuscript and providing helpful
suggestions to improve this paper.
Ellisembia pseudokaradkensis sp. nov. (China) ... 817
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 819-829
https://doi.org/10.5248/132.819
Colpoma guadueticola sp. nov. in a guadual forest
from Quindio Department, Colombia
TANIA RAYMUNDO!, RICARDO SOTO-AGUDELO? & RICARDO VALENZUELA!
"Instituto Politécnico Nacional, Escuela Nacional de Ciencias Bioldgicas,
Plan de Ayala y Carpio s/n Col. Santo Tomas,
Ciudad de México, CP 11340, México
? Docente del Programa de Biologia, Universidad del Quindio,
Armenia, Departamento del Quindio, Colombia
* CORRESPONDENCE TO: rvalenzg@ipn.mx
ABSTRACT—Colpoma guadueticola is described as a new species growing on fallen culms
of Guadua angustifolia from Colombia. The specimens were collected in a guadual forest
from the Natural Reserve “La Montafia del Ocaso, municipality of Quimbaya, Quindio
Department, Colombia. A key to the species of Colpoma is presented.
Key worps—Leotiomycetes, Rhytismatales, Rhytismataceae, taxonomy
Introduction
Colpoma was described by Wallroth in 1833 with C. quercinum (Pers.) Wallr.
as the type species (Darker 1967). The ascomycete genus is characterized by
its elongated (often curved) or ellipsoid apothecial ascomata that erupt under
periderm or break through naked wood and open with a single longitudinal
slit, well developed thick dark colored covering and basal layers, abundant
slender paraphyses that often coil or bend at the tip, and continuous cylindric
or filiform ascospores (Darker 1967, Johnston 1991). Quélet placed Colpoma
within Patellariaceae (Hysteriales), a family later transferred to Phacidiales by
Saccardo (Theon 1918). Darker (1967) moved Colpoma to Hypodermataceae
(Phacidiales) after which Sherwood (1979) and Johnston (1991) transferred the
genus to Rhytismataceae (Phacidiales). Currently, Medardi (2002, 2004) and
Kirk et al. (2008) include Rhytismataceae within Rhytismatales.
820 ... Raymundo, Soto-Agudelo & Valenzuela
The Natural Reserve ‘La Montafia del Ocaso’ is located at 4°34’08”N
75°51'03”W, south of Quimbaya municipality in Quindio Department,
Colombia (Agudelo & Gémez 2001). The guadual forest of this natural
reserve is a floristic resource of great ecological importance because of
the huge diversity of plants, animals, and fungi. The forest is dominated
by Guadua angustifolia Kunth (Poaceae), known as “Colombian bamboo”
and considered the world’s third largest bamboo, reaching heights up to 30
meters (Agudelo & Gémez 2001, Moreno et al. 2007).
There are few studies on fungal species associated with Guadua angustifolia
in Colombia. Restrepo-F. et al. (2005) isolated four agaricomycete genera
on Guadua: Pleurotus spp., Schizophyllum commune Fr., Daedaleopsis
tricolor (Bull.) Bondartsev & Singer, and Polyporus sp. Soto-Agudelo et al.
(2016a) described 29 ascomycete species collected in a guadual forest from
the Natural Reserve ‘La Montafia del Ocaso’ from Colombia; of these, 15
grew on decayed debris of Guadua angustifolia, ten were found on decayed
wood of trees, and four were parasites of other ascomycetes. In 2015 and
2016, during an ongoing survey of Ascomycetes in Colombia (Soto-Agudelo
et al. 2016a,b), three Colpoma specimens were gathered from Guadua
angustifolia. These collections had a distinct morphotype that could not
be satisfactorily accommodated in any of the known species of this genus.
The objective of the present work is to describe these specimens as a new
species of Colpoma associated with Guadua angustifolia in the Department
of Quindio in Colombia.
Materials & methods
The specimens were collected in The Natural Reserve ‘La Montana del Ocaso’in
October 2015 and May 2016 and are deposited in the herbarium of the University
of Quindio, Armenia, Colombia (HUQ) and in the fungal collection ‘Dr. Gaston
Guzman Huerta at the Herbarium Escuela Nacional de Ciencias Bioldgicas of Instituto
Politécnico Nacional, Mexico City, Mexico (ENCB). Latitude/longitude coordinates
were obtained with GPS eTrex (Garmin). Colors are coded according to Kornerup
& Wanscher (1978). Morphological examinations were conducted using protocols
outlined by Johnston (1991) and Minter (1997). Anatomical characters were measured
from rehydrated tissues in 5% aqueous KOH and Melzer’s reagent. Macroscopic
characters were photographed with a Nikon D7000 and micrographs were taken with a
Nikon Coolpix 9000. The meanings of some terms are based on Ulloa & Hanlin (2012)
dictionary.
Colpoma guadueticola sp. nov. (Colombia) ... 821
Fics 1-4: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 1. Gregarious linear
apothecia on fallen culm. 2. Mature fusiform, curved, sigmoid, sinuous to undulate apothecia
with rounded or acute ends. 3. Elongated apothecium with rounded ends. 4. Apothecia ellipsoid,
elongated, lacrymoid with rounded or acute ends. Scale bars: 1 = 10 mm; 2 = 3 mm; 3,4= 1mm.
822 ... Raymundo, Soto-Agudelo & Valenzuela
Taxonomy
Colpoma guadueticola Raymundo, Soto-Agudelo & R. Valenz., sp. nov. Fics 1-15
MycoBank MB 820664
Differs from Colpoma quercinum by its long, filiform ascospores, its paraphyses
dichotomically to irregularly branched at the apex, and its substrate of fallen culms of
Guadua angustifolia.
Type: Colombia, Quindio department: municipality of Quimbaya, Natural Reserve ‘La
Montafia del Ocaso’, 4°34’08”N 75°5103”'W, alt. 1050 m, 29 October 2015, T. Raymundo
5898 (Holotype, HUQ; isotype, ENCB).
EryMo.ocy: Referring to Guaduetum, the phytosociological name for the guadual
forest dominated by Guadua angustifolia, where the specimens were collected.
APOTHECIA (1—)2-10 mm long and 0.4—0.8 mm broad, stromatic, erumpent,
deeply immersed within substratum, and aligned with the main axis of the
fallen culm; visible initially as small ellipsoid to oblong or linear black spots
when closed, later opening by one longitudinal split, variable in shape when
mature: ellipsoid, oblong, fusiform, curved, sigmoid, lacrymoid, sinuous to
undulate, rounded or acute at both ends; the apothecial wall mostly covered
with black stromatic tissue. HyMEeNrum slightly raising the substratum
surface, yellowish white (4A2) to pale yellow (4A3) in young specimens
or at the edges and ends of the ascomata, light yellow (4A5) to deep yellow
(4A8) in mature fresh specimens. STROMATIC TISSUE in vertical transverse
section covering the lateral sides and the lower part of the apothecia,
lateral walls 60-100 um thick, forming a textura angularis, dark brown to
black in KOH, comprising mostly angular cells, olivaceous green to black,
slightly thick-walled 5-9 um diam.; lower wall 30-40 um thick, forming a
textura angularis, dark brown in KOH, comprising mostly angular cells,
olivaceous green to brown, slightly thick-walled, 5-7 um diam. MEDULLAR
EXCIPULUM 30-50 um thick, forming a textura intricata with interwoven
hyphae, hyaline in KOH, septate, 1-2 um diam. SUBHYMENIUM 8-12 um
thick, forming a textura intricata, interwoven hyphae hyaline in KOH, 1-2
um diam. PARAPHYSES filiform, hyaline in KOH, non-septate, smooth, thin
walled, dichotomously to irregularly branched at the apex, extending up to
20 um beyond the asci, 1-2 um diam. Ascr 136-180 x 5-7 um, cylindrical,
Fics 5-11: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 5. Apothecium, vertical
transverse section. 6. Hymenium. 7. Medullary excipulum, hymenium, and stromatic tissue.
8. Lateral stromatic tissue with textura angularis. 9. Basal stromatic tissue. 10-11. Basal and lateral
stromatic tissue and medullary excipulum. Scale bars: 5 = 300 um; 6 = 80 um; 7 = 100 um; 8 = 40
um; 9, 10 = 50 um; 11 = 20 um.
823
Colpoma guadueticola sp. nov. (Colombia) ...
A i tl
> = .
ar, Ah, |
yA
ae |
824 ... Raymundo, Soto-Agudelo & Valenzuela
8-spored, hyaline in KOH, uniformly thin-walled, with only one wall layer
visible, rounded apex, inamyloid. AscosPorgs parallel to subparallel within
the ascus, (68—)80-110(-120) x 0.8-1.0 um, filiform, straight to slightly
curved, hyaline in KOH, aseptate, thin-walled, smooth, gelatinous sheath
not observed at the apex.
ECOLOGY & DISTRIBUTION—Gregarious on fallen and dead culms of
Guadua angustifolia in guadual forest mixed with tropical dry forest. The
species is known only from the type locality.
ADDITIONAL SPECIMENS EXAMINED—COLOMBIA, QUINDIO DEPARTMENT:
Municipality of Quimbaya, Natural Reserve ‘La Montafia del Ocaso, 4°34’08”N
75°51'03”W, alt. 1050 m, 29 October 2015, R. Soto-Agudelo 60 (HUQ); 17 May 2016,
T. Raymundo 6036 (HUQ).
COMMENTS—Colpoma guadueticola is characterized by its variably shaped
apothecia with pointed or rounded ends, yellow hymenium, paraphyses
that are dichotomously to irregularly branched at the apex, long straight to
slightly curved aseptate ascospores, and its host Guadua angustifolia.
Colpoma azaleae (Schwein.) Cooke, C. morbidum (Peck) Sacc., C. quercinum,
and C. styriacum Remler resemble C. guadueticola in having ascospores
longer than 70 um, but C. azaleae is separated by its small (1-3 mm
long) apothecium, dark subrufescent hymenium, and Rhododendron
periclymenoides (Ericaceae) substrate; C. morbidum is distinguished by its
white to gray hymenial surface and growth on Picea mariana (Pinaceae) and
Chamaecyparis thyoides (Cupressaceae); C. quercinum has roughly circular
to oblong apothecia, apices of the paraphyses developing curls and spirals,
and growth on Quercus spp.; and C. styriacum is separated by its small
oblong yellowish green apothecia, 7—8-septate ascospores, and growth on
Vaccinium myrtilloides (Ericaceae). The other Colpoma species are separated
by their shorter (<75 um) ascospores and different hosts, as shown in the
key below.
Key to the species of Colpoma
1. Ascospores ellipsoid to oblong-elliptical, 5-10 um diam., 0-3-septate ............ 2
1. Ascospores filiform, 1-3 um, diam., 0-8-septate .......... 0. eee eee ee eee eee 3
2. On fallen bark of Agathis australis
(apothecia 0.6-1 x 0.3-0.5 mm, broad ellipsoid to oval, hymenial surface
Figs 12-15: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 12. Paraphyses
dichotomously to irregularly branched at the apex. 13. Asci and paraphyses. 14. Asci with parallel
to subparallel ascospores. 15. Ascospores. Scale bars: 12, 14 = 30 um; 13, 15 = 25 um.
Colpoma guadueticola sp. nov. (Colombia) ... 825
’
i )
826 ... Raymundo, Soto-Agudelo & Valenzuela
yellow to orange, ascospores 17-25 x 5-7.5 um, ellipsoid, (0-)1 septate;
known only from New Zealand; Johnston 1991: 405) ................ C. agathidis
2. On fallen wood of Nothofagus
(apothecia 1.5-5 x 1-2.5 mm, ellipsoid to irregular in shape,
hymenial surface translucent yellow, ascospores 31-46 x 6-10 jum, ellipsoid,
(0-)3-septate; known only from New Zealand; Johnston 1991: 407) .. C. nothofagi
St ASCO SP OLS x7 OmLIUNY LON dees Fou bine ea hoot Seer tcndey he cody fre cosy Peepndesake-w ody nse Mee toga 4
Be USGS POLES 7.55 NNN els acihgs Me cihgs OE ech OO ccs ed och Ad edz Ad eds Md bem AS Pegs a Me beget a 14
ALTOVA OM ICL Se teen e ee cee caches fedlagcah ger SUMP es PN pr Te STP ri ARS gor AEE pr TASS pr Sly 5
He CUNEAMOTOSP ORS. . Norn cc Winns © Bees Wien « Whowates « Mowat: Mines « Miri” eben oy Reval Comat 10
5. On Cupressaceae
(on living and dead branches of Juniperus communis, apothecia
1-3 x 1-2 mm, circular, ellipsoid to irregular in shape, hymenial surface
pale gray to fawn, ascospores 40-50 x 1-2 um, almost straight, aseptate;
Buropeanspecios: Miner LO96ae-Ly me iP ncc28F bo 82 bag os eg bo AB Be C. juniperi
DA ANAC GE Rs 2 co Saad usted tes hes den shud’ s cap tude woh Ses ap ok Em pees Doe ees abet be 6
ON
. Paraphyses unbranched, straight, slightly swollen at the apex
(apothecia small, 0.6-1 x 0.28-0.4 mm, long ellipsoid, round or acute at
each end, hymenial surface gray to dark gray, ascospores 33-53 x 2.5-3.5 um,
fili-fusiform, 3-septate at maturity; known only from China on dead or
senescent twigs of Pinus armandii; Hou & Piepenbring 2005: 360) .. C. intermedium
nN
. Paraphyses unbranched or branched, not swollen at the apex
(circinate or flexuous with hooked tips, apothecia larger than 1] mm) ........... 7
7. Apothecia with crenulate margin
(1-2 x 0.5-0.75 mm, ellipsoid, subglobose to irregularly oblong and flexuous,
hymenial surface grayish blue to violaceus gray, ascospores 70-75 x 2 um,
almost straight, rounded apex and basal end acute, aseptate; known only
from Sweden on fallen twigs of Pinus sylvestris; Saccardo 1891: 1128) ..C. serrulatum
7. Apothecia with smooth, striate or rugose margin .............. 0... e eee eee eee 8
8. Paraphyses branched, circinate
(apothecia 1-4 x 0.7-1 mm, ellipsoid to linear, hymenial surface pale
yellowish gray, ascospores 30-45 x 1.5-2 um, aseptate; known only
from USA on decorticated conifer wood; Sherwood 1979: 35)........ C. deustum
8. Paraphyses unbranched, flexuous with hooked tips ...................00 00005. 9
9. Apothecia 0.7-3 x 0.4-1 mm, sometimes curved or irregularly branched,
hymenial surface grayish to bluish gray (ascospores 26-42 x 1.5-2 um,
rod-shaped to filiform, aseptate; European species, growing on fallen twigs
Ol Picegau es siiiapet O97 s-Wiie ote sates Jc tes oe Le Mey, os te C. crispum
9. Apothecia 2-3 mm long, oblong to linear, flexuous, black,
(ascospores 25-35 x 1-1.5 um, subfiliform, rounded apex and acute base,
aseptate; known only from Sweden, growing on bark of fallen twigs
ot Picee dbiessSaccatdo: 1891 1 QY So RP tide Paige! C. pseudographioides
10.
10.
li
1
—_—
12:
12:
Ls
h3y
14.
14.
TS.
15.
16
Colpoma guadueticola sp. nov. (Colombia) ... 827
Apothecia with milk white hymenial surface
(ascospores 50-75 um long, filiform; known only from North America
on dead branches of Rhododendron groenlandicum; Peck 1876: 69,
SAG Cava oul a O04 et, fete Witinte wo ate We aM eaten. tatein' tag ekumetgeee A C. lacteum
Apothecia with gray, bluish gray or grayish blue hymenial surface ............ 11
Paraphyses unbranched, straight to flexuous, apex not differentiated,
apotheatasmial ly <oUsS ti OM aes sae ona Ate + easd Ne egah dhe somaey oe veoh hese aes ea 12
. Paraphyses unbranched or branched, coiled or convoluted at the apex,
21010) Unt sli eve F6 ND N telly aa mde Ramee ey ae eed ae Malem eee tee Ss Wea geil.) Sedat ced 13
Apothecia, 0.7-1.2 x 0.4-0.6 mm ellipsoid to almost circular
(paraphyses straight, ascospores 35-62 x 2-3 um, fili-fusiform, tapering
towards the base, aseptate; known only from China on dead or senescent twigs
of Rhododendron sp.; Hou & Piepenbring 2005: 363) ................ C. sinense
Apothecia 0.5-1.5 x 0.25-0.35 mm, oblong
(paraphyses straight to flexuous, ascospores 35-70 x 1.5-2 um, filiform,
straight to slightly curved, aseptate; known only from China on dead branches
of Rosa sp.; Trotter 1972: 458 as Lophodermium rosae Teng) ............ C. rosae
Apothecia small (0.5-5 x 0.4-1 mm)
(elliptical to oblong elliptical, aligned with the main axis of the substratum,
ascospores 33-55 x 1-2 um, almost straight, aseptate; European species,
on dead attached twigs of Rhododendron tomentosum; Minter 1996b: 1) ...... C. ledi
Apothecia much larger (30-35 x 2-3 mm)
(irregularly oblong, tubular, curved, sigmoid, sometimes also with bifurcations,
ascospores 60-75 x 1-1.5 um, slightly curved, aseptate; known only from Italy
on dead attached twigs of Castanea; Medardi 2004: 36) .............. C. caesium
On dead wood of conifers,
(apothecia ellipsoid, oblong to orbicular, white to gray in hymenial surface,
ascospores 75-100 x 1.5 um, filiform, only known from boreal America on
Picea mariana and Chamaecyparis thyoides; Saccardo 1883: 803, Ellis & Everhart
1892: 723 as Clithris morbida (Pk.) Ellis & Everhart) .............. C. morbidum
OMAN BHOSPETINS iat 5 Se ese gine o: atl des. wag! Pas wag Epp Sed! Eo paul Te bed Ze peed Empe bee deny 15
Paraphyses dichotomously to irregularly branched at the apex
(apothecia 2-10 x 0.4-0.8 mm, ellipsoid, oblong, fusiform, curved, sigmoid,
lacrymoid, sinuous to undulate, round or acute at both ends, hymenial surface
light yellow to deep yellow, on fallen and dead culms of Guadua angustifolia
Ba te Pola (ps Pe sats dm va Male nk Pe 2 Sh C. guadueticola
Paraphyses unbranched, circinate or coiled at the apex ...................008 16
. On Fagaceae and living/dead branches of Quercus spp.
(apothecia 2-15 x 0.5-3 mm, fusiform, navicular to curved, flexuous,
hymenial surface yellow, greenish yellow to olivaceous yellow, ascospores
70-95 x 1-3 um, slightly curved, multiseptate, paraphyses curled to coiled,
hooked; European species; Saccardo 1883: 722, Dennis 1981: 226) .... C. quercinum
PALS SPIGA CCRE RH; Da Sets reach sapt eon sed ale es gh ate gb ate gE mea gli sac cc ESR ec SO 17
828 ... Raymundo, Soto-Agudelo & Valenzuela
17. On dead bark of Rhododendron periclymenoides
(apothecia 1-3 mm long, linear to elongate, flexuous, hymenial surface
dark red, ascospores 90 x 2 um, filiform, aseptate).
Known only from boreal America (Cooke 1889: 58, Saccardo 1891: 1128,
Ellis 8-Evyerharte 1892: /22-as- CHR riIsye Pin) Vs oe Een tao es C. azaleae
17. On fallen dead twigs of Vaccinium myrtilloides
(apothecia 1-2 x 0.6-1 mm, elongate, hymenial surface translucent yellowish
green, ascospores 64-91 x 2-3 um, sigmoid, attenuated base, 7-8 septate).
Known only from Austria (Remler 1980: 173) ...............0.. C. styriacum
Acknowledgments
We wish to express our gratitude to Dr. Peter R. Johnston and Dr. Martin Esqueda
for their useful comments and manuscript review. The authors thank the authorities
of Natural Reserve “La montafia del Ocaso” and University of Quindio for the support
and facilities to carry out the present study. Tania Raymundo thanks IPN for financial
research support in the projects SIP-20161166 and SIP-20170846. Ricardo Valenzuela
thanks COFAA and IPN for financial research support in the projects SIP-20161164
and SIP-20170845.
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 831-837
https://doi.org/10.5248/132.831
Baeomyces lotiformis sp. nov. from China
SHUNAN CAO’, JIANFENG HE’, FANG ZHANG’, HUIMIN TIAN’,
CHUANPENG Livu?, HAIYING WANG** & QIMING ZHOU?*
' Key Laboratory for Polar Science SOA, Polar Research Institute of China,
No. 451 JinQiao Road, Pudong District, Shanghai 200136, China
? Medical Faculty of Chifeng University,
No. 1 YingBin Road, Hongshan District, Chifeng, Inner Mongolia 024000, China
° School of Life Science and Technology, Harbin Institute of Technology,
2 Yikuang Street, Harbin 150001, China
* Key Laboratory of Plant Stress Research, College of Life Science, Shandong Normal University,
No. 88 Wenhua Road, Jinan 250014, China
*CORRESPONDENCE TO: * biowhy@sina.com ° genbank@vip.sina.com
ABSTRACT—A new species of lichenized fungus, Baeomyces lotiformis, is described and
illustrated. It is distinguished by its short podetia and wide apothecium discs, and its affinity
was confirmed by ITS sequence analysis.
KEY worps—Ascomycota, Baeomycetaceae, chemistry, molecular systematics, morphology
Introduction
The worldwide lichen genus Baeomyces Pers. is known from eight crustose
to squamulose species with dark red to brownish stiped apothecia containing
stictic acid as a major secondary substance (Asahina 1943, Upreti 1985, Gierl
& Kalb 1993, Ihlen 1997, Aptroot & Seaward 1999, Kirk et al. 2008). In Yunnan
we encountered an undescribed species that showed affinities with this genus
but deviated conspicuously by its confluent lotus-like apothecia on very short
stipes. Here we present our analysis of the morphology, anatomy, chemistry,
and phylogeny of this species to clarify its taxonomy.
Materials & methods
The material used in this study was collected in 2012 in Yunnan province, China. The
collections examined are preserved in the Lichen Section of the Botanical Herbarium,
832 ... Cao & al.
TABLE 1. nrDNA ITS sequences used in the phylogenetic analysis
SPECIES LOCALITY GENBANK No.
Ainoa mooreana Sweden KJ462262
Japan KJ462263
Anamylopsora pulcherrima Buryatia Republic, Russia KRO17064
— AF274089
Baeomyces lotiformis Yunnan, China KT597690
Baeomyces placophyllus China DQ001274
Xizang, China KT601493
Baeomyces rufus Yunnan, China KT601494
France AF448458
Parainoa subconcolor Yunnan, China KRO17133
Phyllobaeis crustacea Hainan, China KC414617
Hainan, China KC414618
Phyllobaeis imbricata Ecuador KC414619
— HQ650635
Placopsis contortuplicata Antarctica DQ534479
Antarctica KT601492
Placopsis macrophthalma Kerguelen AY212820
Trapelia placodioides — KT695380
Trapelia thieleana — KU672616
Trapeliopsis flexuosa — HQ650634
Trapeliopsis glaucolepidea — AY600082
*New sequences are marked in bold font; missing data are indicated with “—”.
Shandong Normal University, Jinan, China (SDNU). A Zerss Axioskop 2 plus
compound microscope and Motic SMZ-168 dissecting microscope were used to study
the anatomy and morphology. A 10% KOH solution, a 5% NaOCl bleaching solution,
concentrated alcoholic p-phenylenediamine (PD), Lugol’s Iodine solution, and thin-
layer chromatography (TLC) (Culberson & Kristinsson 1970, Culberson 1972, White &
James 1985) were used to detect lichen substances.
DNA extraction, PCR amplification, and sequencing
Total DNA was extracted using a modified CTAB method (Cao et al. 2013).
PCR amplifications were performed using a Biometra T-Gradient thermal cycler.
The nrDNA ITS region was amplified by the primer pair ITS5 and ITS4 (White et al.
1990). The 50-ul reaction volume comprised 1 ul total DNA, 2 ul of each primer (10
uM), 1 ul Taq polymerase (rTaq DNA Polymerase, 5 U/ul), 4 wl dNTP (2.5 mM each),
5 ul amplification buffer (10x, 25 mM MgCl, contained), and 35 ul ddH,O. Cycling
parameters included initial denaturation at 95°C for 5 min, 30 cycles of denaturation
at 94°C for 40 s, annealing at 52°C for 40 s, extension at 72°C for 2 min, and a final
extension at 72°C for 10 min. A negative control, without DNA template, was prepared
in every amplification series so as to minimize the possibility of contamination.
Baeomyces lotiformis sp. nov. (China) ... 833
PCR products were purified using a Biocolor gel purification kit (BioScience &
Technology Co. Ltd.) and sequenced using an ABI 3730 XL DNA Sequencer.
Four nrDNA ITS sequences, including one from the new species, were obtained by the
authors, and another 17 nrDNA ITS sequences representing six species were downloaded
from GenBank (TaBLE 1). Three sequences from Placopsis (Trapeliaceae), two from
Trapelia (Trapeliaceae) and two from Trapeliopsis (Trapeliaceae) were selected as outgroup.
Phylogenetic analysis
The sequences were aligned using ClustalW 1.6 (Thompson et al. 1994). The
phylogenetic analysis was executed with software Mega 7 (Kumar et al. 2016). The
Kimura-2-parameter was selected as the nucleotide substitution model, and gaps or
missing data were set as pairwise deletion. The neighbor-joining (NJ) method was used
in constructing the phylogenetic tree and the reliability of the inferred tree was tested by
1000 bootstrap replications.
Result and discussion
The NJ-tree (Fic. 1) of the ITS rDNA sequences clearly supports the genera
Baeomyces, Phyllobaeis, Parainoa, Ainoa, and Anamylopsora in Baeomycetaceae
as sister branches. The bootstrap values were 98% for Baeomyces and 97% for
Phyllobaeis. Baeomyces lotiformis clustered within the Baeomyces branch, but
was clearly separated from B. placophyllus and B. rufus.
100 r— Baeomyces placophylluse DQ001274
99 Baeomyces placophyllus KT601493
Baeomyces rufus AF448458
100 Baeomyces rufus KT601494
Baeomyces lotiformis [HT] K1T597690
100 Phyllobaeis imbricata HQ650635
Phyllobaeis imbricata KC414618
97 Phyllobaeis crustacea HQ650835
100! Phyllobaeis crustacea KC414617
98
saoAwoseg
99
66
syaeqolAud
56
aeaaRPjaoAwoseg
Parainoa subconcolor KRO17133
Ainoa mooreana KJ462263
100 Ainoa mooreana KJ462262
99
Anamylopsora pulcherrima KR017064
100! Anamylopsora pulcherrima AF274089
100 Placopsis contortuplicata KT601492
100 Placopsis contortuplicata DQ534479
Placopsis macrophthalma AY 212820
90 Trapeliopsis glaucolepidea AY600082
Trapeliopsis flexuosa HQ650634
63 Trapelia thieleana KU672616
100 Trapelia placodioides KT695380
aeaoeyedes
0.020
Fic. 1. Neighbor-joining tree of the nrDNA ITS region sequences. Nucleotide: K2 model, gaps or
missing data were partial deletion, bootstrap = 1000. Genetic distance scale = 0.02. Numbers at
nodes present the bootstrap support value (numbers <50 not shown). New sequences are set in
bold font; holotype sequence indicated by ‘[HT]-
834 ... Cao & al.
Taxonomy
Baeomyces lotiformis S.N. Cao, sp. nov. Fi. 2
MycoBank MB 819009
Differs from the other Baeomyces species by its confluent, lotus-like apothecia and
crustose thallus.
Type: China, Yunnan: Jianchuan County, 26°23’N 100°49.8’E, alt. 2665 m, on soil, 25
Nov. 2012, H. Y. Wang yn013 (Holotype, SDNU 20160915; GenBank KT597690).
ErymMo oey: Latin lotiformis, referring to the lotus-like apothecia. The Latin noun
“lotus” has the compounding form “lot-”; and the sufhx indicating a morphological
similarity is “-formis”.
THALLUS crustose, gray green to ochre, tightly attached to the substrate, lacking
a cortical layer, irregularly delimited; surface irregular with many small,
mammillary warts, 0.2 mm in diameter; algae layer continuous, algal cells
green, ovoid or ellipsoid, 9-11 x 10-13 um.
APOTHECIA dark red to reddish brown, scattered over the thallus, when
full-grown with very sinuous margins deeply dividing the disc, which is
thus seemingly composed of many confluent discs aggregated into a lotus-
like structure, about 2 mm in diameter, when young rounded and 0.5 mm in
diameter; disc flattened with swollen margin; podetia very short, less than 1
mm, with nearly sessile apothecia, lacking algae; hymenium 162.5-175 um
thick, algae not observed within subhymenium and exciple, I-; paraphyses
branched, not septate; asci long-clavate, 8-spored, with apex I-, 75-90 x 7.5
uum; ascospores oblong or fusiform, hyaline, simple, 8-10 x 5 um (10 spores
observed).
CHEMISTRY—Spot tests: Thallus K+ yellow, C-. Specimen contains stictic
acid (TLC).
ADDITIONAL MATERIAL EXAMINED: CHINA, YUNNAN: Mt. Ailao, 24°21’N 102°31.2’E,
alt. 2300 m, on rock, 25 November 2010, H.Y. Wang yn076 (SDNU 20160916).
CoMMENTSs: Baeomyces lotiformis is characterized by a crustose thallus and
dark red to reddish brown apothecia with very short podetia, which show
affinities with the well-known stiped crustose to squamulose lichen genera
Baeomyces, Dibaeis Clements, and Phyllobaeis Kalb & Gierl. However, the
new species differs chemically from Dibaeis and Phyllobaeis: stictic acid is
the main substance, and baeomycesic and norstictic acid are absent. The
morphological and chemical characters support inclusion of B. lotiformis
in Baeomyces, but within this genus its confluent, lotus-like apothecia are
unique.
Baeomyces lotiformis sp. nov. (China) ... 835
Fic. 2. Baeomyces lotiformis (holotype, SDNU 20160915). a, habit; b, apothecia; c, granular
mammillary warts (in water); d, asci.
In view of its very short podetia, we compared B. lotiformis with the
crustose lichen genera also currently included in Baeomycetaceae: Ainoa
Lumbsch & I. Schmitt, Anamylopsora Timdal, and Parainoa Resl & T. Sprib.
(Liicking et al. 2016). Ainoa differs by its dark-brown exciple, narrowly
ellipsoid ascospores (16-32 6-14 um), and by the presence of gyrophoric
acid restricted to apothecia and pycnidia (Brodo & Lendemer 2015). Parainoa
is similar to Ainoa but differs in containing depsidones (like Baeomyces);
however, Parainoa completely lacks a differentiated and extended hypothecial
stalk, its hypothecium is yellowish, its thallus is creamish white, its ascomata
may or may not contain gyrophoric acid, and it is characterized by simple,
roundish apothecia (Resl et al. 2015). Anamylopsora is still more distinct by
836 ... Cao & al.
its squamulose thallus, its black sessile apothecia, its subspherical ascospores,
and the presence of alectorialic acid (Timdal 1991).
Our ITS rDNA sequence analysis clusters B. lotiformis with B. rufus and
B. placophyllus on a well-supported branch and support B. lotiformis as a new
species in the genus Baeomyces.
Acknowledgments
Our thanks are due in the first place to the Chinese Arctic and Antarctic
Administration (2011GW12016). Our project was financially supported by the
Program for Scientific Research Innovation Team in Colleges and Universities of
Shandong Province, the National Natural Science Foundation of China (31270059),
the Science Foundation of Jinan (201202024), and the National innovation Fund
for college students (201310445082). We thank Prof. Jae-Seoun Hur, Prof. Sergey
Kondratyuk, and Dr. Harrie Sipman for presubmission reviews. We also thank Dr.
Shaun Pennycook for valuable comments and nomenclature reviews.
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Cao SN, Wei XL, Zhou QM, Wei JC. 2013. Phyllobaeis crustacea sp. nov. from China.
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Gierl C, Kalb K. 1993. Die Flechtengattung Dibaeis. Eine Ubersicht iiber die rosafriichtigen Arten
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MJ, Redhead S, Schoch CL, Spatafora JW, Stalpers JA, Vilgalys R, Aime MC, Aptroot
A, Bauer R, Begerow D, Benny GL, Castlebury LA, Crous PW, Dai YC, Gams W, Geiser
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Kirk PM, Cannon PF, Minter DW, Stalpers JA. 2008. Dictionary of the fungi. 10th edition.
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 839-847
https://doi.org/10.5248/132.839
Lirula sichuanensis sp. nov.
on Picea likiangensis var. rubescens from Sichuan, China
*
CHUN-LIN YANG’, XIU-LAN XU’”, ZHENG-GAO ZHANG: & YING-GAO LIU?
‘College of Forestry, Sichuan Agricultural University,
Huimin Road 211, Chengdu, Sichuan 611130, China
? Forestry Research Institute, Chengdu Academy of Agriculture and Forestry Sciences,
Nongke Road 200, Chengdu, Sichuan 611130, China
> Huangshan Yunle Ganoderma Co., Ltd. of Anhui Province,
Yingkan Road, Jingde, Anhui 242600, China
* CORRESPONDENCE TO: lyg927@263.net
ABSTRACT—Lirula sichuanensis on 2-year-old dead needles attached on living twigs of Picea
likiangensis var. rubescens [= P. likiangensis var. balfouriana] in Sichuan Province is described
and illustrated as a new species. Phylogenetic analysis based on the internal transcribed
spacer sequences of ribosomal DNA sequences confirms its distinction from similar species
of the genus. The type specimen is deposited in the Forest Protection Laboratory of Sichuan
Agricultural University, China (SAUF).
Key worps—Rhytismataceae, needle blight, taxonomy, ITS rDNA
Introduction
Lirula Darker is distinguished from other genera of Rhytismataceae by
its nervisequious ascomata and ascospore shape (Darker 1967). Initially, six
species were recombined in Lirula by Darker (1967): L. abietis-concoloris
(Mayr) Darker, L. macrospora (R. Hartig) Darker, L. mirabilis (Darker) Darker,
L. nervata (Darker) Darker, L. nervisequa (DC.) Darker, and L. punctata
(Darker) Darker. Ziller (1969) described a new species, L. brevispora Ziller
on Picea glauca (Moench) Voss from Canada. Kaneko (1993) described
L. pakistanensis S. Kaneko on Abies pindrow (Royle ex D. Don) Royle from
Pakistan and later (Kaneko 2003) described L. exigua S. Kaneko and L. japonica
840 ... Yang & al.
S. Kaneko on Abies mariesii Mast. from Japan. Fan et al. (2012) described
L. yunnanensis L. Fan et al. on Abies georgei Orr [= A. forrestii var. georgei (Orr)
Farjon] from Yunnan Province, China.
The eleven Lirula species usually cause severe needle blight on conifers; two
species parasitize Picea, and the others mainly Abies (TaBLE 1). In this paper,
we report a new species of Lirula on dead needles of Picea likiangensis var.
rubescens [= P. likiangensis var. balfouriana (Rehder & E.H. Wilson) Hillier].
Materials & methods
Mature fruitbodies were selected and examined macroscopically under a dissecting
microscope at 6-45x magnification and microscopically under a compound microscope
at 40-100x magnification (objective lens). The fruitbodies were sectioned by hand;
vertical sections were mounted in water for observation of ascomatal and conidiomatal
outlines. Gelatinous sheaths surrounding ascospores and paraphyses were examined in
water or cotton blue lactophenol. The colours of anatomical structures and ascospore
contents were observed in water. All measurements were made using material mounted
in water.
Total genomic DNA was extracted from ascomata following the operations manual
of DNA extraction kit™ (Tiangen, China). The internal transcribed spacer sequences
of ribosomal DNA (ITS rDNA) region were amplified with PCR using the primers
ITS1/ITS4 (White et al. 1990). PCR was performed in 50-uL reactions including DNA
template 5 uL, primer ITS1/ITS4 2 wL, 2x MasterMix 25 uL, and ddH,O 16 uL, under
the following parameters: 95 °C for 1 min., 50 °C for 1 min., 72 °C for 1 min., for a total
TABLE 1. Hosts of Lirula species.
SPECIES Host REFERENCE
Darker 1967; Scharpf 1988
Ziller 1969
Kaneko 2003; Fan et al. 2012
L. abietis-concoloris Abies concolor
L. brevispora Picea glauca
L. exigua Abies mariesii, A. georgei
L. japonica
L. macrospora
L. mirabilis
L. nervata
L. nervisequa
L. pakistanensis
L. punctata
L. yunnanensis
Abies mariesii
Picea likiangensis var. rubescens, P. wilsonii,
Picea asperata, P. crassifolia, P. jezoensis,
P. koraiensis, P. neoveitchii, P. schrenkiana,
P. abies, P. glauca, P. pungens
Abies balsamea
Abies balsamea
Abies alba, A. fabri, A. fargesii, A. holophylla,
A. forrestii var. smithii, Taxus wallichiana
Abies pindrow
Abies amabilis, A. balsamea, A. concolor, A.
grandis, A. procera, A. mariesii
Abies georgei
Kaneko 2003; Fan et al. 2012
Darker 1932; Darker 1967;
Lin et al. 2012; Scharpf 1988
Darker 1967
Darker 1967
Darker 1967; Lin et al. 2012;
Fan et al. 2012
Kaneko 1993
Darker 1967
Fan et al. 2012
Lirula sichuanensis sp. nov. (China) ... 841
of 35 cycles followed by a final extension step at 72 °C for 5 min. The PCR products
detected by gel electrophoresis were sent to Invitrogen Biotechnology (Guangzhou,
China) for purification and sequencing.
The ITS rDNA sequences were aligned with ClustalW 1.8 (Thompson et al. 1997);
sections of the sequences longer than the sequence of new species were excluded from
the analysis. A neighbor-joining (NJ) tree was generated with MEGA5.1 software
(Tamura et al. 2011), and branch stability was estimated with 1000 bootstrap pseudo-
replicates calculated with Kimura's two-parameter model (Kimura 1980). Other ITS
rDNA sequences used in this study were downloaded from GenBank (TaBLE 2). The
newly generated sequences were deposited in GenBank.
TABLE 2. ITS rDNA sequences of the species used in the study and their GenBank
accession numbers. The holotype of L. sichuanensis is indicated as [HT].
SPECIES VOUCHER GENBANK No. REFERENCE
Lirula macrospora AF462440 NCBI
AF462441 NCBI
Lirula yunnanensis HQ902156 Fan et al. 2012
Lirula exigua HQ902157 Fan et al. 2012
HQ902154 Fan et al. 2012
HQ902155 Fan et al. 2012
Lirula sichuanensis SAUF1605001 [HT] KY399845 This paper
SAUF1606001 KY399846 This paper
Lophodermium conigenum AY422489 Pan et al. 2006
Lophodermium australe KM106810 Oono et al. 2014
KM106812 Oono et al. 2014
Lophodermium indianum KF636510 NCBI
KF636502 NCBI
Rhytisma panamense GQ253102 Hou et al. 2010
Rhizosphaera kalkhofhi KM435342 Azeem et al. 2015
JQ353721 You et al. 2013
Taxonomy
Lirula sichuanensis X.L. Xu, C.L. Yang & Y.G. Liu, sp. nov. FIGs 1-12
MycoBank MB 819592
Differs from Lirula macrospora by its larger asci and ascospores, its branched, unswollen
(or only slightly swollen) paraphyses, and its hypophyllous conidiomata forming a
continuous or interrupted row on both sides of the substrate needle’s stomatal line.
Type: China, Sichuan Province, Pingwu County, Yangdonghe Plantation, alt. 2822 m, on
needle of Picea likiangensis var. rubescens Rehder & E.H. Wilson (Pinaceae), 5 May 2016,
X.L. Xu & C.L. Yang 160504 (Holotype, SAUF1605001; GenBank KY399845).
ETtyMoLoGy: sichuanensis, referring to Sichuan Province where the specimens were
collected.
842 ... Yang & al.
AscoMaTA hypophyllous (occasionally epiphyllous), nervisequious, on
previous year dead needles but still attached to a living twig; dull to shining black
in surface view, (2-—)4-12(-17) x 0.38-0.6 mm, no perimeter line, variably long
and extending at times almost the entire length of the needle; intra-epidermal in
median vertical section, 190-330 um deep; strongly raised above the surface of
the substrate, opening widely by a single longitudinal split, exposing the canary
yellow hymenium. COVERING STROMA conspicuous and well carbonized, no
lip cell observed; <96-133 um thick near the ascomal centre, becoming thinner
towards the edges and not extending to the basal layer; consisting of an outer
layer of host cuticle and epidermal cell remnants, an inner layer of brown to
black brown textura angularis with 6-11 um diam. cells, and an innermost
layer of colourless to pale brown textura globulosa with 7-12 um diam. cells.
HYMENIUM concave to flat, (80-)100-210 um thick. SUBHYMENIUM hyaline,
deep concave, composed of hyaline, pseudoparenchymatous cells. BASAL
STROMA poorly developed or almost absent. ParApuHysES filiform, straight or
curved, longer than asci, 180-240 um long, 2.6-4.2 um thick at the base, multi-
septate, tapering towards the tip, 1.4—2.3 um thick, not or only slightly swollen,
often branched near the top or in the middle, surrounded by a gelatinous matrix
1.7-2.7 um thick, but not forming an epithecium. Asc clavate, subtruncate at
the apex when maturing, tapered below to a short stipe, 105-166 x 16-23 um,
J-, 8-spored. Ascosporss cylindrical-clavate, hyaline, arranged in fascicles,
51-90 x 5-7 um, rounded at the apex, tapered towards the base, encased in a
conspicuous (3.7-6.6 um thick) hyaline gelatinous sheath.
Conip1oMaTA hypophyllous, on both sides of the stomatal lines, forming
two continuous or interrupted rows on dead needles attached to living twigs,
110-360 um wide, intra-epidermal in vertical section, 58-65 um deep;
concolorous with the substrate or pale brown, appearing as shining blisters,
round or ellipse or irregular shape, opening by sparse little ostioles. UPPER
WALL absent. BASAL WALL extremely poorly developed. Conip1a hyaline,
baculiform, straight or slightly curved, 4-7 x 2 um.
ZONE LINES not observed.
HABITAT & DISTRIBUTION: Producing conidiomata and ascomata on dead
needles still attached to living twigs. Known only from Sichuan Province,
China.
Fics. 1-8. Lirula sichuanensis (holotype, SAUF1605001) sexual stage on Picea needles. 1. Ascomata
observed under the dissecting microscope. 2. Ascoma in vertical section. 3. Covering stroma near
the centre of ascoma in vertical section. 4. Asci and paraphyses. 5. Paraphysis bases and a young
ascus. 6. Paraphysis tips. 7. Ascus containing eight ascospores. 8. A single released ascospore.
843
Lirula sichuanensis sp. nov. (China) ...
844 ... Yang & al.
stomata
pycnidia
Figs. 9-12. Lirula sichuanensis (SAUF1606001) asexual stage on Picea needles. 9. Pycnidia
observed under the dissecting microscope. 10. Pycnidium in vertical section. 11. Sporogenous
tissue. 12. Conidia.
ADDITIONAL SPECIMENS EXAMINED—CHINA, SICHUAN PROVINCE, Pingwu County,
Yangdonghe Plantation, alt. 2822 m, on Picea likiangensis var. rubescens, 13 June
2016, X.L. Xu & C.L. Yang 160613 (SAUF1606001; GenBank KY399846); GANzI
AUTONOMOUS PREFECTURE, Luhuo County, alt. 3240 m, on Picea likiangensis var.
rubescens, 30 July 2014, X.L. Xu & C.L. Yang 140730 (SAUF1407001).
Discussion
Few gene sequences of Lirula are available in GenBank. ITS rDNA sequence
analysis clustered L. sichuanensis in an independent lineage with L. macrospora
and L. yunnanensis ona branch with 70% bootstrap support apart from L. exigua
in a distant clade. Lirula exigua differs from L. sichuanensis, L. macrospora, and
L. yunnanensis (Darker 1932, Fan et al. 2012) in its paraphyses not swollen at
tips, narrower asci, shorter ascospores, and conidiomata that are scattered and
not in continuous rows. Additionally, L. exigua has been observed in needles
on dead twigs; the areas of needles with ascomata are whitened and fragile;
and the ascomal color and shape are somewhat different (Fan et al. 2012).
Lirula sichuanensis sp. nov. (China) ... 845
100, Lophodermium indianum KF636510
Lophodermium indianum KF 636502
Lophodermium australe KM106810
ea ae australe KM106812
Lophodermium conigenum AY 422489
Rhytisma panamense GQ253 102
100, Lirila sichuanensis KY399845
Lirula sichuanensis iY 399846
Lirula yunnanensis HQ902156
100, Lirula macrospora AF462440
Lirula macrospora AF 462441
100/-Lirula exigua HQ902154
Lirula exigua HQ902155
93 / | Lirula exiguaHQ902157
100 -Rhizosphaera kalkhoffii JQ353721
Rhizosphaera kalkhoffii KM435342
99
86
0.05
|
Fic. 13. Neighbor-joining phylogeny generated from partial ITS sequence analysis of Lirula
sichuanensis and related species, using Rhizosphaera kalkhoffii as outgroup. Bootstrap values
>80% from 1000 replications are shown on the respective branches.
Consequently, L. exigua needs further taxonomic study (including comparison
with the type species, L. nervisequa) to establish whether it belongs in Lirula.
Lirula sichuanensis differs from other species in its distinctive paraphyses,
which are longer and thicker, multi-septate, usually curved, and not to only
slightly swollen at the tips (often branched near the middle or the apex) and
thicker gelatinous sheath. In addition, the ascospores and gelatinous sheath are
thicker, and the conidia are broader. The new species shares a similar habit
with L. brevispora and L. macrospora on the needles of Picea spp. (Darker 1932,
Ziller 1969) but differs from them by its longer thicker branched paraphyses,
conspicuously larger or wider ascospores and gelatinous sheath, and
inconspicuous basal stroma. Lirula macrospora differs in its smaller (100-132
x 14-16 um) clavate-cylindric asci, much smaller (56-68 x 2.5-3 um) clavate
ascospores, unbranched paraphyses with swollen tips, irregularly scattered or
sometimes 2-3 coalescent conidiomata that do not form two continuous or
interrupted rows on needles (Darker 1932), much smaller (2.5-4.5 x 1.2-1.5
um) conidia, and sometimes producing trichogynes (Lin et al. 2012). Ascal
shape and size of L. sichuanensis is similar to that of L. yunnanensis, which
differs by its bluish grey to bluish black ascomata, its narrower ascospores often
846 ... Yang & al.
covered by gelatinous caps and a thinner gelatinous sheath, and its bigger,
deeper, pale yellowish brown, epiphyllous, nervisequious pycnidia (Fan et al.
2012).
Lirula sichuanensis causes serious needle blight of Picea likiangensis var.
rubescens in planted forests in Sichuan. It usually infects 1-year-old needles of
20- to 30-year-old trees from May to July and in the following March to May
produces ascomata on 2-year-old needles. According to our survey, this disease
spreads quickly in planted spruce forests. An overall investigation is needed
to understand the scope and area affected by the disease, and this pathogen
should be controlled at ecological security level by forestry measures or drug
treatment.
Acknowledgments
The authors are grateful to Prof. Y.R. Lin and Dr. H.D. Zheng for serving as pre-
submission reviewers leading to the improvement of our manuscript and to Dr. Shaun
Pennycook for critical review of the manuscript and nomenclature help. This study was
supported by the Forest Pest Management and Quarantine Station of Sichuan Province
for the Ecological Forest Diseases and Pests Control along the Upper Yangtze River.
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Darker GD. 1932. The Hypodermataceae of conifers. Published by the Arnold Arboretum of
Harvard University 1. 131 p.
Darker GD. 1967. A revision of the genera of the Hypodermataceae. Canadian Journal of Botany
45(8): 1399-1444. https://doi.org/10.1139/b67-145
Fan L, Lin YR, Wang S, Hou CL. 2012. Two species of Lirula on Abies from Yunnan, Southwest
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Hou CL, Trampe T, Piepenbring M. 2010. A new species of Rhytisma causes tar spot on
Comarostaphylis arbutoides (Ericaceae) in Panama. Mycopathologia 169(3): 225-229.
https://doi.org/10.1007/s11046-009-9250-4
Kaneko S. 1993. Parasitic fungi on woody plants from Pakistan Himalaya. Cryptogamic Flora of
Pakistan 2(2): 149-168.
Kaneko S. 2003. Two new species of Lirula on Abies from Japan. Mycoscience 44(4): 335-338.
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Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through
comparative studies of nucleotide sequences. Journal of Molecular Evolution 16(2): 111-120.
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Lin YR, Liu HY, Hou CL, Wang SJ, Ye M, Huang CL, Xiang Y, Yu SM. 2012. Flora fungorum
sinicorum. vol. 40, Rhytismatales [in Chinese]. Science Press, Beijing.
Oono R, Lutzoni FE, Arnold AE, Kaye L, U’ren JM, May G, Carbone I. 2014. Genetic variation
in horizontally transmitted fungal endophytes of pine needles reveals population structure
in cryptic species. American Journal of Botany 101(8): 1362-1374.
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Pan X, Liu YG, Zou LK, Peng PH, Chen WD, Liu YG. 2006. Cloning and sequencing of the ITS
regions of Lophodermium spp. Biotechnology 16(4): 5-7.
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https://doi.org/10.1139/b69-037
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 849-855
https://doi.org/10.5248/132.849
Hypoderma rubi on two new hosts in Slovakia
MARTIN PASTIRGAK! & KATARiNA PASTIRCAKOVA”
‘National Agricultural and Food Centre, Plant Production Research Institute,
Bratislavskd 122, SK-92168 Piestany, Slovakia
*Slovak Academy of Sciences, Institute of Forest Ecology, Branch for Woody Plant Biology,
Akademickd 2, SK-94901 Nitra, Slovakia
“ CORRESPONDENCE TO: uefezima@hotmail.com
ABSTRACT—Hypoderma rubi was found on the previous year’s fallen petioles of Fraxinus
chinensis subsp. rhynchophylla and on attached dead twigs of Rhododendron fortunei in
Slovakia. The fungus, which is recorded for the first time on these host taxa, also represents a
new taxon for the Slovak mycota. The morphological characteristics of the fungus found on
Slovak collections are described.
Key worps—Chinese ash, Fortune's rhododendron, morphology, Rhytismataceae,
Rhytismatales
Introduction
Hypoderma rubi, the type and best-known species of Hypoderma
(Rhytismataceae, Rhytismatales), occurs on a large number of herbaceous
and woody plant genera and has an extensive geographical distribution.
According to Farr & Rossman (2017), most records are from Asia (China,
India), although the fungus has also been recorded in New Zealand, North
America (USA), Central America (Panama), and Europe (Czech Republic,
Germany, Spain, Sweden, Ukraine, United Kingdom). European findings of
the fungus are mainly on the type host genus Rubus L. (Hilitzer 1929, Lotz-
Winter et al. 2011, Moreno et al. 2004, Eriksson 2014, Lantz et al. 2011,
Dudka et al. 2004, Cannon et al. 1985). There are also some specimens of
H. rubi from other European countries (Channel Islands, Italy, Norway),
Asia (Japan), South America (Chile), and Africa (Malawi) in the K & IMI
850 ... Pastiréak & Pastircakova
fungaria (Royal Botanic Gardens, Kew) that have not yet been formally
published.
In 2015, during a survey of the mycota of trees and shrubs in Mlynany
Arboretum of the Slovak Academy of Sciences, two collections of H. rubi
were made. This species is not listed in the checklist of fungi of Slovakia
(Lizon & Bacigalova 1998), and there are no previous records of it in Slovakia.
Petrak’s records of H. rubi on Rubus spp. (deposited in international herbaria,
e.g. BPI, IMI, K) collected in the former Czechoslovakia, originate in
the territory of Czech Republic. This paper presents, therefore, the first
collections of H. rubi from Slovakia, and the first on Fraxinus chinensis
subsp. rhynchophylla (Hance) A.E. Murray and Rhododendron fortunei
Lindl. A description and illustrations of the fungus based on the Slovak
collections are provided.
Materials & methods
The previous year’s petioles of FE. chinensis subsp. rhynchophylla and attached
dead twigs of R. fortunei were collected in the Mlyhany Arboretum (south-west of
Slovakia) in autumn 2015. The specimens were examined using an Olympus SZ61
stereomicroscope and Olympus BX51 standard light microscope. Morphological
and microscopic examinations were carried out on dried material rehydrated in
water. Lactophenol blue solution (Merck, Darmstadt) was used to stain hyaline
structures. The morphological structures of the fungus were photographed using an
Olympus SP350 digital camera. We compared our measurements with previously
published descriptions. Voucher specimens are deposited in the Plant Pathology
Herbarium of the Institute of Forest Ecology of Slovak Academy of Sciences, Nitra,
Slovakia (NR).
Taxonomy
Hypoderma rubi (Pers.) DC. ex Chevall.,
J. Phys. Chim. Hist. Nat. Arts. 94: 31 (1822) FIGURE 1
= Hypoderma virgultorum DC., Fl. Frang., Ed. 3, 6: 165 (1815), nom. illeg.
= Hypoderma commune (Fr.) Duby, Mém. Soc. Phys. Hist. Nat. Genéve 16: 53 (1861)
HyYSTEROTHECIA embedded in the previous year’s fallen Chinese ash
petioles and in paler portions of Fortune’s rhododendron twigs, abundant,
subcuticular, black, elliptical to elongated-elliptical, raising the substrate
surface, (0.4—)0.8-2.7 x 0.3-0.6 mm, opening by a single longitudinal split,
lips grey, clypeus thickened towards the lips. Lip cELLs hyaline, 14-20.5 x
2.5-3.5 um. Asci unitunicate, clavate, long-stalked, (75-)90-125 x
10-14.5 um, 8-spored. Ascospores hyaline, aseptate, guttulate, elongated-
New hosts for Hypoderma rubi (Slovakia) ... 851
FiGuRE 1. Hypoderma rubi (A, C-H = NR 5477; B, I-L = NR 5478). A: Hysterothecia, pycnidia
and zone lines on Fraxinus petiole; B: Hysterothecia on Rhododendron twig; C: Mature
hysterothecium (surface view); D: Hysterothecium in vertical section; E: Lip cells; F: Asci with
ascospores; G: Paraphyses; H: Ascospores; I: Pycnidia (surface view); J: Pycnidium (detail of
surface appearance); K: Conidiogenous cells with conidia; L: Conidia. Scale bars: A, B = 1 mm;
C=0.5 mm; D = 100 um; E, H, K =5 um; F G= 10 um; I = 200 um; J = 50 um; L = 2 um.
clavate to cylindrical, bent towards the top end, 18-25 x 2.5-3.5 um, usually
not covered by a gelatinous sheath but sometimes clad in a thin sheath.
852 ... Pastiréak & Pastircakova
TABLE 1. Biometric characteristics of Hypoderma rubi from the Slovak material
and as reported by other authors.
MoRPHOLOGY CANNON & MINTER (1986) JOHNSTON (1990) SLOVAK MATERIAL
Hysterothecia (mm) 1.4-2.5(-3.0) x 0.5-0.75 0.8-3.0 x 0.3-0.6 (0.4-)0.8-2.7 x 0.3-0.6
Asci (um) 87-130 x 12-14.5 110-160 x 11-14 (75-)90-125 x 10-14.5
Ascospores (um) (16—)20-24(-30) x 2.5-3.5 15-28 x 2.5-5.5 18-25 x 2.5-3.5
Pycnidia (um diam.) 75-250(-300) 100-300 (80-)105-220
Conidia (um) 2-4 x 0.75-1 3-5 x 1-1.5 2-4x1
PARAPHYSES hyaline, filiform, septate, branched or unbranched, not swollen
at the apex, as long as mature asci, covered by a gelatinous sheath. PYCNIDIA
subcuticular, black, circular, (80-)105-220 um diam., opening by 1-2
small central ostioles, accompanying the hysterothecia. CONIDIOGENOUS
CELLS cylindrical, tapering towards the apex, hyaline, 7-12 x 1-2 um.
ConipiIA cylindrical, sometimes allantoid, hyaline, aseptate, 2-4 x 1 um.
ZONE LINES black, thin.
SPECIMENS EXAMINED—SLOVAKIA. ViEsKA NAD ZiITAvou, Mlynhany Arboretum,
48.3203°N 18.3668°E, on the previous year’s petioles of Fraxinus chinensis subsp.
rhynchophylla (Oleaceae), 2 Oct. 2015, leg. et det. K. Pastir¢éakova (NR 5477);
48.3208°N 18.3692°E, on dead attached twigs of Rhododendron fortunei (Ericaceae),
9 Nov. 2015, leg. et det. M. Pastircak (NR 5478).
The fungus was identified as Hypoderma rubi based on morphological
description and measurements of reproductive structures. A comparison of
some biometric characteristics of H. rubi on Rubus sp. (Cannon & Minter
1986), on different host plants Johnston 1990), and of the Slovak material is
given in TABLE 1. Morphometric data of the Slovak collections correspond
well to previously published descriptions. Although ascospores usually are
not covered by a gelatinous sheath, a thin sheath was often observed after
staining (Fic. 1H). Hou et al. (2007) also recorded H. rubi on Cunninghamia
lanceolata (Lamb.) Hook. (Cupressaceae) with ascospores clad in a thin
gelatinous sheath.
The findings of H. rubi on F. chinensis subsp. rhynchophylla and R. fortunei
from Slovakia represent the first records of the fungus in the country and
new host taxa for the fungus.
Discussion
Rubus is the principal host genus on which Hypoderma rubi is most
commonly collected (Minter 1984). However, according to Farr & Rossman
New hosts for Hypoderma rubi (Slovakia) ... 853
(2017), H. rubi has been recorded on leaves, stems, and petioles of species
representing 35 host genera in 27 families. The species therefore seems to
be widely plurivorous, although Cannon & Minter (1986), who doubt the
conspecificity of the fungi on these hosts, suggest that numerous taxa may
be involved. According to Lantieri et al. (2011), the genetic relationships
amongst specimens identified as H. rubi are poorly resolved and the
application of this name by different authors is uncertain. There are as yet
very few molecular data available for H. rubi and from only a few hosts
(Lantieri et al. 2011, Lantz et al. 2011). A comprehensive phylogenetic study
is necessary to determine their identity and relationships.
Hypoderma rubi on Rhododendron sp. has been recorded in Europe
(Wales; Smith 1951) and Asia (Malaysia; Spooner 1991). In China,
four other Hypoderma species are known on Rhododendron spp.:
H. rhododendri-mariesii Y.R. Lin & S.J. Wang on R. mariesii Hemsl. & E.H.
Wilson (Lin et al. 2004), H. cuspidatum C.L. Hou & M. Piepenbr., H. shiqii
C.L. Hou & M. Piepenbr., and H. urniforme C.L. Hou & M. Piepenbr. on
Rhododendron sp. (Hou & Piepenbring 2006, Hou et al. 2007). They differ
from H. rubi mainly by the shape and position of their hysterothecia as well
as ascospore characteristics. Hypoderma rhododendri-mariesii has smaller
(<860 um long) hysterothecia, asci with cylindric-fusiform ascospores,
and shorter ((9—)11-17 um) ascospores; H. cuspidatum has intraepidermal
hysterothecia with acute ends, ascospores with thick gelatinous sheaths, and
unbranched paraphyses; H. shiqii has intraepidermal hysterothecia without
lips that often open with one slit and additional lateral fissures, clavate asci
with bifusiform ascospores, and ascospores with thick gelatinous sheaths;
and H. urniforme is characterised by urniform, intraepidermal hysterothecia
with red-brown lips and almost cylindrical asci with ellipsoidal to long-
ellipsoidal ascospores with irregular gelatinous sheaths. H. cuspidatum,
H. shigii, and H. urniforme often occur together on the same twigs (Hou et
al. 2007).
On-line databases of the British Mycological Society (Fungal Records
Database of Britain and Ireland, http://www.fieldmycology.net) and Royal
Botanic Gardens, Kew (Herb. IMI database, http://www.herbimi.info)
include data on fungarium specimens of H. rubi on dead fallen leaves and
dead attached twigs of Rhododendron ponticum L., R. sinogrande Balf. f. &
W.W. Sm. and R. sp. from Scotland (IMI 358124b, 358144, 358155, 358157,
358158a); and on dead fallen petioles of Fraxinus excelsior L. from England
(IMI 339025, K(M)160355). These records are evidently not yet published.
854 ... Pastiréak & Pastircakova
In the USA, Hanlin (1963) recorded Hypoderma commune on Fraxinus
americana L.; however Powell (1974) treated H. commune as a synonym
of H. rubi. Based on the available literature and electronic resources, we
consider that only one Hypoderma species, H. rubi, has been recorded on
Fraxinus.
Acknowledgments
We sincerely thank Roger T.A. Cook (UK) and Brian M. Spooner (UK) for kindly
correcting the English and for their valuable comments, which helped to improve the
manuscript. The authors are also indebted to Peter R. Johnston (Landcare Research,
Auckland, New Zealand) and Hyeon-Dong Shin (Korea University, Seoul, South
Korea) for critical comments on the manuscript and for acting as peer-reviewers.
This study was supported by the Scientific Grant Agency of the Ministry of Education
of the Slovak Republic and of Slovak Academy of Sciences, projects VEGA No.
2/0071/14 and No. 2/0183/14.
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(Ascomycota) from China. Nova Hedwigia 84: 487-493.
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Mycotaxon 118: 393-401. https://doi.org/10.5248/118.393
Lantz H, Johnston PR, Park D, Minter DW. 2011. Molecular phylogeny reveals a core clade of
Rhytismatales. Mycologia 103: 57-74. https://doi.org/10.3852/10-060
New hosts for Hypoderma rubi (Slovakia) ... 855
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the Botanical Garden of the University of Frankfurt. Zeitschrift fiir Mykologie 77: 89-122.
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Parque Nacional de Cabanferos (Ciudad Real). Boletin de la Sociedad Micoldgica de Madrid
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Powell PE. 1974. Taxonomic studies in the genus Hypoderma (Rhytismataceae). PhD dissertation,
Cornell University, Ithaca.
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Spooner BM. 1991. Lophodermium and Hypoderma (Rhytismatales) from Mt. Kinabalu, Sabah.
Kew Bulletin 46(1): 73-100. https://doi.org/10.2307/4110745
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 857-865
https://doi.org/10.5248/132.857
New reports of Myriospora (Acarosporaceae) from Europe
KERRY KNUDSEN’, JANA KOCOURKOVA' & ULF SCHIEFELBEIN”
" Czech University of Life Sciences Prague, Faculty of Environmental Sciences, Department of Ecology,
Kamycka 129, Praha 6 - Suchdol, CZ-165 21, Czech Republic
? Bliicherstrafse 71, D- 18055 Rostock, Germany
* CORRESPONDENCE TO: knudsen@fzp.czu.cz
ABsTRACT—Myriospora dilatata is newly reported for the Czech Republic and M. myochroa
new for Italy. Myriospora rufescens was rediscovered in Germany almost 100 years after
its first collection. A neotype is designated for Acarospora fusca, which is recognized as a
synonym of M. rufescens.
Key worps—Mpyriospora hassei, Silobia, Trimmatothelopsis
Introduction
The genus Myriospora in the Acarosporaceae is a well-supported clade
distinguished by a constellation of morphological characters (non-lecideine
apothecia, high hymenium, thin paraphyses, interrupted algal layer, short
conidia, no secondary metabolites or norstictic acid) (Wedin et al. 2009;
Westberg et al. 2011, 2015). The genus currently contains 12 species that
occur in Antarctica, Asia, Europe, and North and South America (Knudsen
2011, Westberg et al. 2011, Knudsen et al. 2012, Knudsen & Bungartz 2014,
Schiefelbein et al. 2015, Purvis et al. in press). Myriospora fulvoviridula (Harm.)
Cl. Roux is a synonym of M. scabrida (H. Magn.) K. Knudsen & Arcadia
(Knudsen et al. 2017, Roux et al. 2014). The most common species in the genus
is M. smaragdula (Wahlenb.) Nageli ex Uloth, which occurs in Asia, Europe,
North and South America (Magnusson 1929, Knudsen 2007, Westberg et al.
2011, Knudsen et al. 2012). The center of diversity of the genus appears to be
in Europe where nine species occur. The genus was originally treated as Silobia
858 ... Knudsen, Kocourkova & Schiefelbein
(Westberg et al. 2011, Knudsen 2011) and briefly as Trimmatothelopsis [(Roux
& Navarro-Rosinés 2011); for a current circumscription of Trimmatothelopsis
see Knudsen & Lendemer 2016]. It is now treated under its oldest available
name Myriospora (Arcadia & Knudsen 2012).
While the reported diversity of Acarosporaceae is high in Europe (Magnusson
1929, 1956), no species is particularly frequent except for Acarospora fuscata
(Schrad.) Arnold. It is not unusual to collect all day on rocks and soil in the
Czech Republic, Germany, or the Italian Alps and only find one or two species
of Acarosporaceae besides A. fuscata. Because of this diversity of taxa and
infrequency of populations, not to mention problems identifying many species,
there are possibilities for new discoveries. In this paper, we report three new
records of Myriospora species in Europe.
Materials & methods
Myriospora specimens from institutional herbaria (PRM, STU, TBS, UCR, UPS)
and personal herbaria of Jana Kocourkova and Kerry Knudsen (hb. K & K) and Ulf
Schiefelbein (hb. Schiefelbein) were determined using key and descriptions by Westberg
et al. (2011). They were analyzed with standard microscopy and spot tests (Nash et al.
2002). Undiluted fresh Lugol's solution (Merck) was used to test amyloid reactions of the
hymenium and subhymenium. The description of M. rufescens according to Westberg
et al. (2011) was revised. Jana Kocourkova photographed M. rufescens using a digital
camera Olympus DP72 attached to an Olympus SZX7 Stereomicroscope and Olympus
BX51 fitted with a Nomarski differential interference contrast and image stacks
processed with software Olympus Deep Focus 3.1.
Taxonomy
Myriospora dilatata (M. Westb. & Wedin) K. Knudsen & Arcadia,
Opusc. Philolich. 11:21, 2012.
= Silobia dilatata M. Westb. & Wedin, Lichenologist 43(1): 12, 2011.
Type: Sweden. Torne Lappmark. Jukkasjarvi par., Tornehamn, shore of Lake
Tornetrask, on silicate rock, 3 Sept. 2006, E. Baloch SW116 (S [n.v.], holotype).
=Trimmatothelopsis dilatata (M. Westb. & Wedin) Cl. Roux &
Nav.-Ros., Bull. Soc. Linn. Provence 62: 176, 2011.
DESCRIPTION. See Westberg et al. (2011, as Silobia dilatata). For pictures see
Westberg et al. (2011) and Wirth et al. (2013).
EcoLocy & DISTRIBUTION. Ferrophilous, heavy metal tolerant, often in
moist habitats near streams. Czech Republic (Bohemia), Germany (Wirth et
al. 2011, 2013), Norway (Westberg et al. 2011), Sweden (Westberg et al. 2011).
SPECIMEN EXAMINED. CZECH REPUBLIC. NorRTHERN BOHEMIA. Krkonoése Mts,
Snézka Mt., SSE of former mine ‘Kovarna, 50°43’35”N 15°43’58.7’E, alt. 1075 m, humid
New Myriospora reports for Europe ... 859
lower edge of boulder scree, mica-schist outcrop enriched of heavy metals (copper ores
and arsenopyrite), 8 Sept. 2016, J. Kocourkova 9083 & K. Knudsen (hb. K & K).
Discussion. Myriospora dilatata is easily identified among Myriospora by
its orange to reddish areoles, non-punctiform apothecia, lack of secondary
metabolites, and occurrence on substrates rich in iron and other heavy metals
(Westberg et al. 2011). The species is reported as rare in Bohemia in the Czech
Republic. In the KrkonosSe Mountains M. dilatata was growing on a mica-
schist outcrop enriched with heavy metals (copper ores and arsenopyrite) in a
relatively moist habitat with abandoned mining pits. In these same mountains,
the often red-powdered ferrophilous species Acarospora sinopica (Wahlenb.)
Korb. is common (hb. K & K) and there have been several collections by
Zdenék Palice (PRA) and V. Kutak (PRM) of the orange ferrophilous species
M. tangerina (M. Westb. & Wedin) K. Knudsen & Arcadia (Westberg et
al. 2011, Knudsen et al. 2017). Punctiform apothecia easily distinguish
A. sinopica from M. dilatata with its areoles with dilated apothecia. With
difficult specimens the full description and discussion of A. sinopica in
Magnusson (1929) is informative. Punctiform apothecia also distinguish
M. tangerina from M. dilatata as well as large squamules. An apparently cryptic
ferrophilous taxon of Myriospora, similar in appearance to M. dilatata and
sometimes identified as Acarospora smaragdula f. subochracea H. Magn., was
reported in an unpublished molecular analysis of specimens from the Alps
but not yet described (Westberg et al. 2011). This report extends the range of
M. dilatata from Fennoscandia to Central Europe.
Myriospora myochroa (M. Westb.) K. Knudsen & Arcadia,
Opusc. Philolich. 11:22, 2012.
= Silobia myochroa M. Westb., Lichenologist 43(1): 14, 2011.
Type: Sweden. Bohuslan. Orust, Morlanda Parish, near car park at Stocken, grid ref:
(RT90) 6456258 1241653, c. 1 m, on the vertical surface of a SW-facing rock, 16 Sept.
2003, A. Crewe & O. W. Purvis 719 (S [n.v.], holotype).
= Trimmatothelopsis myochroa (M. Westb.) Cl. Roux & Nav.-
Ros., Bull. Soc. Linn. Provence 62: 176, 2011.
DESCRIPTION. For description and pictures Westberg et al. (2011, as Silobia
myochroa). For picture and map of Italian distribution, see Nimis & Martellos
(2017).
ECOLOGY & DISTRIBUTION. Asia, Europe. Often on silicate rock underhangs
at low to high elevations. In salt spray zone along Atlantic coast. Czech Republic
(Malicek & Vondrak 2016), Finland (Westberg et al. 2011), France (Roux et
al. 2014), Germany (Wirth et al. 2013, Schiefelbein et al. 2015), Italy, Norway
860 ... Knudsen, Kocourkova & Schiefelbein
(Westberg et al. 2011), Poland (Flakus 2014), Russia (Siberia) (Zhdanov 2013),
Sweden (Westberg et al. 2011).
SELECTED SPECIMEN EXAMINED. ITALY. FRIULI. Giogaia dei Fleors: sen tiero vs la
cima, 2406 m, in silicate rock underhang, 17 Aug. 1996, M. Tretiach 24565 (TBS).
Discussion. Our discovery of specimens of Myriospora myochroa in TBS
extends the species range into Italy. If spot tests are negative and crystals of
norstictic acid are not observed in the cortex with polarized light, M. myochroa
is sometimes misidentified as M. hassei (Herre) K. Knudsen & Arcadia, a
species we currently consider endemic to the Pacific Plate Lichen Bioregion on
the south and central coast of California (Knudsen 2011, Knudsen & Bungartz
2014, Nimis 2016, Nimis & Martellos 2017).
Myriospora rufescens (Ach.) Hepp ex Uloth, Flora 44: 618, 1861. FIG. 1
= Sagedia rufescens Ach., Lichenogr. Univ. 329, 1810.
TYPE: Great Britain. England. Turner 23 (H-ACH-980[n.v.]), lectotype (designated
in Westberg et al. 2011).
= Lecanora rufescens (Ach.) Nyl., Flora 55: 364, 1872.
= Acarospora rufescens (Ach.) Kremp., Lich.-Fl. Bayern.: 173 (1861).
= Silobia rufescens (Ach.) M. Westb. & Wedin, Lichenologist 43(1): 18, 2011.
= Trimmatothelopsis rufescens (Ach.) Cl. Roux & Nav.-
Ros., Bull. Soc. Linn. Provence 62: 176, 2011.
= Acarospora fusca B. de Lesd., Recherch. Lich. Dunkerque 1(Suppl.): 100, 1914, syn. nov.
Type: France. Dunkerque. Malo Terminus, dunes, 1910, B. des Lesdain
s.n. (holotype [n.v.]. Presumed lost in bombing of Dunkirk in WwW2).
Germany. Mecklenburg. West Pomerania, Vorpommern-Greifswald, Greifswald,
Koos Island, eastern edge of the island, 54°10’13”N 13°25’19’E, c. 1 m, on single
boulder on boulder beach, 6 Aug. 2004, U. Schiefelbein 4446 (Neotype designated
here, NY; isoneotype, hb. Schiefelbein).
HyPoTHALLus endosubstratic, I-. THattus epilithic, areolate, areoles
verruciform [areoles with an immersed apothecium dilating until the thallus
is reduced to a thalline margin around the disc and often resembling Lecanora
apothecia and/or areoles not verruciform, rounded to angular, 0.4-0.9(-1.5)
mm diam., on smooth surfaces forming a contiguous indeterminate crust,
becoming dispersed on uneven rock surface, replicating by the division of
the areoles, the upper surface dark grey to dark greyish or reddish brown,
epruinose, smooth to rugulose, flat to somewhat convex, lower surface
broadly attached, epicortical layer lacking, upper cortex (10—)40-60 um thick,
paraplectenchymatous but individual cells difficult to distinguish, <5 um diam.
Algal layer with or without distinct interruption by thin hyphal bundles, ca.
100 um tall, continuous beneath apothecia. Medulla continuous with attaching
New Myriospora reports for Europe ... 861
Fig. 1. Myriospora rufescens (Acarospora fusca neotype, Schiefelbein 4446): A. Dispersed areoles of
the thallus with solitary apothecia; B. Areoles with mostly solitary apothecia forming a contiguous
indeterminate crust; C. Apothecia 1-(2-4) per areole in contiguous crust; D. Vertical section of
ascoma. Scale bars: A~C = 500 um; D = 100 um.
862 ... Knudsen, Kocourkova & Schiefelbein
hyphae, mostly 2-3 um wide, mixed with substrate particles. APOTHECIA
1-(2-4) per areole, immersed, disc rounded, 0.07-0.25(-0.60) mm diam.,
reddish brown to blackening, flat, smooth or rough, epruinose, parathecium
colorless in section, ca. 15 um below the hymenium, expanding up to 60 um
near the surface, uppermost cells with dark brown caps, forming an indistinct
perithecial crown concolorous with the thallus or becoming blacker,
epihymenium reddish brown to dark brown. HyMENtuM (100-)120-140(-160)
tum tall (as low as 85 um per Magnusson 1929), I- red. PARAPHYSES 1-1.5 um
wide at midlevel, sparsely branched, tips cylindrical to clavate, widened to 2.5 um,
asci up to 150 x 27 um. AscosporeEs narrowly ellipsoid to bacilliform, 3-6 x
(1-)1.5(-2) um. SUBHYMENIUM opaque, (20-)40-55 um tall inspersed with oil
drops, I+ blue. Pycnip1A not observed.
ECOLOGY & DISTRIBUTION. Usually on smooth or rough surfaces of silicate
rock. Czech Republic (Magnusson 1929), France (Roux et al. 2014), Germany
(Magnusson 1929, Westberg et al. 2011), Norway (Westberg et al. 2011),
Sweden (Westberg et al. 2011), United Kingdom (Magnusson 1929, Fletcher et
al. 2009, Westberg et al. 2011).
OTHER SPECIMENS EXAMINED. GERMANY. Lower Saxony. District Stade, NefSsand
island: c. 250 m W of the Inselwarthaus, 53°33’20’N 9°45’25’E, c. 5 m, acidophilous,
nutrient-poor grassland with single shrubs, on siliceous stone in the grass, 26 Sept. 2015,
U. Schiefelbein 4415, (hb. Schiefelbein); c. 200 m S of the Inselwarthaus, 53°33’15”N
9°45’40”E, c. 5 m, pile of stone in a nutrient-poor grassland, forming large patch on
siliceous boulder, 26 Sept. 2015, U. Schiefelbein 4416 (hb. Schiefelbein).
Discussion. Myriospora rufescens is distinguished from other members
of the genus by its small (<0.5 mm diam.) apothecia with a relatively short
(usually 120-140 um) hymenium with thin paraphyses and tall subhymenium
inspersed with oil drops (Westberg et al. 2011). As in M. dilatata, the apothecia
are not punctiform but dilated, often forming verruciform areoles reminiscent
of Lecanora apothecia.
Magnusson (1929) considered Acarospora rufescens and A. fusca to
represent distinct species, but we consider them conspecific, representing two
different morphotypes that can intergrade even in the same population. Thus in
Magnusson (1929), A. rufescens was the morphotype with verruciform areoles
often resembling Lecanora apothecia (see Fic. 14 and also lectotype photo in
Westberg et al. 2011) while A. fusca represented the morphotype with mostly
contiguous non-verruciform areoles, often with two or more apothecia and
usually forming a thin indeterminate crust (Fic. 1B,c). The holotype of A. fusca
in Bouly de Lesdain’s herbarium is presumed to have been lost in World War II
during the 1940 bombing of Dunkirk, as were many other specimens cited by
New Myriospora reports for Europe ... 863
= = oe pate
Le Se ae
= eS — > — = : to
Fig. 2. Acarospora fusca neotype locality on boulder beach
in Koos Island, Mecklenburg, Germany.
Magnusson in his Acarospora monograph (Magnusson 1929, Abbeyes 1966).
The only recorded collection of M. rufescens in Germany was made in Anhalt,
Bernburg, by G.H. Zschacke in 1907 (Magnusson 1929, Westberg et al. 2011,
Wirth et al. 2013). This German collection of M. rufescens was identified as
A. fusca (Magnusson 1929). We designate a modern collection by Ulf
Schiefelbein from Koos Island in Germany as the neotype of A. fusca, deposit
it in NY, and formally recognize A. fusca as a synonym of M. rufescens. It has
been over 100 years since M. rufescens was last collected in Germany in 1907.
A return trip to Koos Island revealed no additional M. rufescens on the other
rocks along the shore.
Myriospora rufescens is a small species that can be overlooked, especially
in a mixed saxicolous community. The three 21st century German collections
suggest that M. rufescens prefers microhabitats with high annual relative
humidity. The neotype locality for Acarospora fusca on Koos Island is a boulder
beach on the western edge of Greifswalder Bodden, a bay of the Baltic Sea (Fic.
2). The neotype population occurred on the top of a large boulder that was not
usually inundated but regularly sprayed by waves of brackish water, leaving salt
crystals embedded in the cortex and apothecia. The two other modern German
sites for M. rufescens were on the small narrow island of Nefsand, located just
downstream from Hamburg at the mouth of Elbe River.
864 ... Knudsen, Kocourkova & Schiefelbein
Acknowledgments
We thank our reviewers, J.C. Lendemer (NY) and J. McCarthy, S.J. (Canada). We
thank for their assistance P.L. Nimis (TBS) and the curators of PRA, PRM, STU, TBS,
UCR, and UPS. The work of Jana Kocourkova and Kerry Knudsen was financially
supported by 42900/1312/3166 [Environmental aspects of sustainable development of
society], a grant from the Faculty of Environmental Sciences, Czech University of Life
Sciences Prague.
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 867-874
https://doi.org/10.5248/132.867
Perenniporia puerensis sp. nov. from southern China
WEI-Li Lru’, TaAr-MIN Xv’, SHAN SHEN’,
XIANG-Fu Liu’, YANG SUN' & CHANG-LIN ZHAO**
‘College of Biodiversity Conservation and Utilization, ? College of Life Sciences &
* Key Laboratory of Forest Disaster Warning and Control of Yunnan Province,
Southwest Forestry University, Kunming 650224, P.R. China
* CORRESPONDENCE TO: fungichanglinz@163.com
ABSTRACT—A new polypore, Perenniporia puerensis, collected from Yunnan province,
southern China, is described and illustrated based on four collections using morphology-
based methods. Macroscopically, the new species is characterized by an annual growth
habit, resupinate basidiocarps with a yellow to ochraceous pore surface, and 4-6 pores per
mm. Microscopically, it has a dimitic hyphal system with non-dextrinoid and cyanophilous
skeletal hyphae that are encrusted with pale-yellow crystals, and basidiospores that are ovoid
to subglobose, thick-walled, non-dextrinoid, cyanophilous, and 4.3-5.5 x 3.7-4.7 um.
Key worps— Basidiomycota, Polyporaceae, Polyporales, taxonomy, white rot fungus
Introduction
Perenniporia Murrill is a large, cosmopolitan genus characterized by
poroid basidiomata and basidiospores that are thick-walled, ellipsoid to
distinctly truncate, cyanophilous and variably dextrinoid and amyloid. The
hyphal system in Perenniporia is di- or trimitic with clamp connections on the
generative hyphae and vegetative hyphae that are cyanophilous and variably
dextrinoid or amyloid (Decock & Stalpers 2006). Approximately one hundred
species have been described in or transferred to the genus (Gilbertson &
Ryvarden 1987; Decock & Ryvarden 1999; Hattori & Lee 1999; Decock et
al. 2001; Nuftez & Ryvarden 2001; Choeyklin et al. 2009; Cui & Zhao 2012;
Zhao & Cui 2013a,b; Zhao et al. 2013; Ryvarden & Melo 2014; Jang et al. 2015;
Decock 2016).
868 ... Liu & al.
Phylogenetic studies of Perenniporia s.]. inferred from nuclear ribosomal
LSU and ITS DNA sequence data well support several monophyletic groups
that can be recognized as distinct genera (Robledo et al. 2009; Zhao & Cui
2013a,b; Zhao et al. 2013) within the polyporoid clade.
Fifty Perenniporia species have been recorded during recent taxonomic
surveys in China (Dai et al. 2002, 2015; Dai 2012), including several new
species (Cui et al. 2007; Xiong et al. 2008; Dai 2010; Dai et al. 2011; Cui &
Zhao 2012; Zhao & Cui 2012, 2013a,b; Zhao et al. 2013, 2014). During one
such recent survey, we identified an undescribed species matching the concept
of Perenniporia.
Materials & methods
Cited specimens are deposited at the herbarium of Southwest Forestry University,
Kunming, China (SWFC). Microscopical protocols follow Dai (2012). Sections were
examined at magnifications up to 1000x using a Nikon Eclipse E 80i microscope and
phase contrast illumination. Drawings were made with the aid of a drawing tube.
Microscopic features, measurements, and drawings were made from slide preparations
stained with Cotton Blue and Melzer’s reagent. Spores were measured from sections cut
from the tubes. To present spore size variations, 5% of measurements were excluded
from each end of the range, and extreme values are given in parentheses. In the text
the following abbreviations are used: M = Melzer’s reagent, M- = both inamyloid and
nondextrinoid, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB+ = cyanophilous,
L = mean spore length (arithmetic average of all spores), W = mean spore width
(arithmetic average of all spores), Q = the range of variation in the L/W ratios of n
number of basidiospores, n = number of basidiospores measured from the 4 specimens
studied. Special color terms follow Petersen (1996).
Taxonomy
Perenniporia puerensis C.L. Zhao, sp. nov. Fies 1, 2
MycoBank MB 823635
Differs from Perenniporia straminea by its bigger pores and bigger basidiospores, from
P. tibetica by its smaller pores and the absence of rhizomorphs; and from P. subacida by its
annual growth habit, its non-dextrinoid skeletal hyphae, and its truncate basidiospores.
Type: China. Yunnan Province: Puer, Laiyanghe Nature Reserve, on a fallen angiosperm
trunk, 21 November 2016, CLZhao 606 (Holotype, SWFC 000606).
ErymMo.oey: The specific epithet puerensis (Lat.) refers to the locality (Puer) of the type
specimen.
BASIDIOMATA annual, resupinate, adnate, without odor or taste when fresh,
becoming corky upon drying, <5 x 3 cm, 2.5 mm thick at the center. Pore
surface cream to buff when fresh, yellow to ochraceous upon drying; pores
Perenniporia puerensis sp. nov. (China) ... 869
Wap?
—_ © , Fy? 7
eal a. %
Fic. 1. Perenniporia puerensis (holotype, SWFC 000606): basidiomata. Scale bars = 1 cm.
870 ... Liu & al.
C OC
2O C C
CO @
VV
Cc
a
10 ym
i NON
iN | , ly
) AN'5|!
—_——
7,
A= Pm!
Fi
Cl
— <
\
\
W
:
‘i
\ Z 7
iS
TE
10 um
Fic. 2. Perenniporia puerensis (holotype, SWFC 000606). a. Basidiospores; b. Basidia and basidioles;
c. Cystidioles; d. Hyphae from trama; e. Hyphae from subiculum. Scale bars: a = 5 um; b-e = 10 um.
round to angular, 4-6 per mm; dissepiments thin, entire. Sterile margin
narrow, cream, <1 mm wide. Subiculum cream to buff, thin, <0.5 mm thick.
Tubes concolorous with pore surface, corky, <2 mm long.
HYPHAL STRUCTURE dimitic; generative hyphae with clamp connections;
skeletal hyphae M-, CB+; tissues unchanged in KOH and the pale-yellow
crystals dissolving in KOH.
Perenniporia puerensis sp. nov. (China) ... 871
SUBICULUM generative hyphae infrequent, hyaline, thin-walled, frequently
branched, 2-3.5 um diam.; skeletal hyphae dominant, hyaline, thick-walled
with a wide to narrow lumen, frequently branched, interwoven, 2.5-4.5 um
diam., encrusted with pale-yellow crystals.
TUBE generative hyphae infrequent, hyaline, thin-walled, frequently
branched, 2-3 um diam.; skeletal hyphae dominant, hyaline, thick-walled with
a wide lumen, frequently branched, interwoven, 2-4 um diam., encrusted with
pale-yellow crystals. Cystidia absent, fusoid cystidioles present, hyaline, thin-
walled, 10-16 x 2.5-5 um. Basidia barrel-shaped, with four sterigmata and a
basal clamp connection, 13-18 x 7-10 um; basidioles dominant, mostly pear-
shaped, but slightly smaller than basidia.
BASIDIOSPORES ovoid to subglobose, either truncate or non-truncate,
hyaline, thick-walled, smooth, non-dextrinoid, CB+, (4.1-)4.3-5.5(-5.7) x
(3.5-)3.7-4.7(-5.1) um, L = 4.95 um, W = 4.18 um, Q = 1.14-1.21 (n = 120/4).
TYPE OF ROT: white rot.
ADDITIONAL SPECIMENS EXAMINED: CHINA. YUNNAN PROVINCE. Puer: Laiyanghe
Nature Reserve, on fallen angiosperm trunk, 21 November 2016, CLZhao 607 (SWFC
000607); CLZhao 608 (SWFC 000608); CLZhao 609 (SWFC 000609).
Discussion
Four other Perenniporia species were also found with P. puerensis in the
same locality (Yunnan province, China): P aridula B.K. Cui & C.L. Zhao,
P. bannaensis B.K. Cui & C.L. Zhao, P. piceicola Y.C. Dai, and P. russeimarginata
B.K. Cui & C.L. Zhao. Perenniporia aridula is distinguished by its perennial
basidiomata and larger basidiospores (6-7 x 5.1-6 um, Zhao et al. 2013);
P. bannaensis is distinguished by its smaller pores (6-8 per mm), unbranched
skeletal hyphae, and strongly dextrinoid basidiospores (Zhao et al. 2013);
P. piceicola differs by larger basidiospores (11-14 x 5.4-7.5 um) and the
presence of pyriform cystidia (Dai et al. 2002); and P russeimarginata is
separated by its perennial basidiocarps with a white to cream pore surface
and the distinct reddish brown sterile margin (Zhao & Cui 2013a).
The presence of both truncate and non-truncate basidiospores is reminiscent
of two similar Perenniporia species—P. straminea (Bres.) Ryvarden and P. tibetica
B.K. Cui & C.L. Zhao: P straminea is distinguished by smaller pores (7-9 per
mm) and basidiospores (3-4 x 2.5-3 um; Ryvarden 1988), while P. tibetica
differs in its bigger pores (2-3 per mm) and presence of white to cream-colored
rhizomorphs (Cui & Zhao 2012).
Perenniporia subacida (Peck) Donk, which resembles P. puerensis in pore
size (4-6 per mm) and non-dextrinoid basidiospores, is distinguished by its
872 ... Liu & al.
perennial basidiocarps, strongly dextrinoid skeletal hyphae, and non-truncate
basidiospores (Nufiez & Ryvarden 2001, Ryvarden & Melo 2014).
Polypores are an extensively studied group in Basidiomycota (Gilbertson &
Ryvarden 1987, Nufiez & Ryvarden 2001, Ryvarden & Melo 2014), but Chinese
polypore diversity is still not well known, especially in the subtropics and
tropics where many recently described taxa have been collected (Cui & Dai
2008; Cui et al. 2009, 2010, 2011; Du & Cui 2009; Li & Cui 2010; He & Li 2011;
Jia & Cui 2011; Yu et al. 2013; Yang & He 2014; Chen et al. 2015). The new
species Perenniporia puerensis is also from the subtropics. We anticipate that
additional polypore taxa will be found in China after further investigation and
molecular analyses.
Acknowledgments
Special thanks are due to Dr. Sana Jabeen (University of Education, Pakistan) and
Jason Karakehian (Harvard University, USA) who reviewed the manuscript. We express
our gratitude to Yong-He Li (Yunnan Academy of Biodiversity, Southwest Forestry
University) for his support on molecular work and Kai- Yue Luo, Zhen Xu, and Ting Zeng
(College of Biodiversity Conservation and Utilization, Southwest Forestry University)
for their collection support. The research is supported by the National Natural Science
Foundation of China (Project No. 31700023), and the Science Foundation of Southwest
Forestry University (Project No. 111715) and the Science and Technology Talent
Support Program of Three Areas in Yunnan Province (Project No. 21700329).
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 875-879
https://doi.org/10.5248/132.875
Sporidesmiopsis lushanensis sp. nov.
from Lushan Mountain, China
X1A0-MEI WANG’, ZI-JIAN ZHAO’, SHAN-SHAN CHEN’, XIAO-MAN LIv’,
Hao-Hua LY, X1tu-Guo ZHANG?“ & JI- WEN XIA”®
‘College of Agronomy, Jilin Agricultural University, Changchun, Jilin, 130118, China
? Shandong Provincial Key Laboratory for Biology of Vegetable Diseases and Insect Pests,
College of Plant Protection, Shandong Agricultural University,
Taian, Shandong, 271018, China
* CORRESPONDENCE TO: “sdau613@163.com, * xiajiwen1@126.com
ABSTRACT—Sporidesmiopsis Iushanensis is described and illustrated as a new species from
dead stems of an unidentified broadleaf tree in Jiangxi Province, China. The fungus is
characterized by macronematous conidiophores with apical branches and conidiogenous
cells arising from stipe and branches with 6-7-euseptate conidia. A key to Sporidesmiopsis
species is provided.
Key worps—hyphomycetes, taxonomy, Sordariales
Introduction
Sporidesmiopsis Subram. & Bhat, established in 1989 with S. malabarica
Subram. & Bhat as type species, is distinguished from Sporidesmium
Link by its apically branched mononematous conidiophores and
conidiogenous cells produced on both stipe and branches (Subramanian
& Bhat 1989, Wongsawas et al. 2008). However, S. malabarica was
subsequently determined to be a synonym of the earlier Brachysporiella
dennisii J.L. Crane & Dumont, which was transferred to Sporidesmiopsis
as S. dennisii (J.L. Crane & Dumont) Bhat et al. (Bhat & Kendrick 1993).
Five other species have been described in the genus: S. goanensis Bhat &
W.B. Kendr., S. guangxiensis J.W. Xia & X.G. Zhang, S. malloti J.W. Xia &
X.G. Zhang, S. pluriseptata J.S. Monteiro et al., and S. zhejiangensis Wongs.
876 ... Wang & al.
et al. (Bhat & Kendrick 1993, Wongsawas et al. 2008, Xia et al. 2014, 2015,
Monteiro et al. 2016), but S. zhejiangensis was subsequently transferred
to Ellisembiopsis Santa Izabel & Gusmao, due to its distoseptate conidia
(Santa Izabel et al. 2013). Thus, only five species are currently accepted in
Sporidesmiopsis.
Among our fungal collections from Lushan Mountain, ahyphomycetous
specimen with the morphological characteristics of Sporidesmiopsis was
collected, which we describe here as a new species, S. lushanensis. Type
specimens are conserved in the Herbarium of Department of Plant
Pathology, Shandong Agricultural University, Taian, China (HSAUP) and
Herbarium Mycologium, Institute of Microbiology, Chinese Academy of
Sciences, Beijing, China (HMAS).
Sporidesmiopsis lushanensis J.W. Xia & X.G. Zhang, sp. nov. FIG. 1
MycoBank MB 823681
Differs from Sporidesmiopsis goanensis, S. guangxiensis, and S. malloti by its wider
conidia.
Type: China, Jiangxi Province: Lushan Mountain, on dead stems of an unidentified
broadleaf tree, 25 Oct. 2016, J.W. Xia (Holotype, HSAUP H6725; isotype, HMAS
245644).
ETryMOLoGy: in reference to the type locality.
CoLonigs on the natural substrate effuse, brown to dark brown, hairy.
Mycelium superficial and partly immersed, composed of septate, pale brown,
smooth hyphae, 1-2 um diam. CONIDIOPHORES distinct, single, branched,
erect, straight or slightly flexuous, cylindrical, smooth, thick-walled, brown
to dark brown, 8-15-septate, 250-450 x 10-14 um. CONIDIOGENOUS CELLS
monoblastic, integrated and discrete, cylindrical, truncate at the apex after
conidium secession, 12.5-20 x 5-7 um. Conrp1A solitary, dry, fusiform,
base truncate, apex rounded, brown to dark brown, 30-50 x 11.5-15.5 um,
6—7-euseptate.
CoMMENTS—'hree Sporidesmiopsis species differ from S. lushanensis
by their narrower conidia: S. goanensis (5-7 um; Bhat & Kendrick 1993),
S. guangxiensis (6-7.5 um; Xia et al. 2014), and S. malloti (8.5-11.5 um;
Xia et al. 2015). Sporidesmiopsis dennisii and S. pluriseptata differ from
S. lushanensis by their longer conidia with much more numerous septa (see
Fic. 2).
Sporidesmiopsis lushanensis sp. nov. (China) ... 877
Fic. 1. Sporidesmiopsis lushanensis (holotype, HSAUP H6725).
A, B. Conidiophores, conidiogenous cells, and conidia; C. Conidia. Scale bars = 20 um.
878 ... Wang & al.
A
a a as a
EPL Ee
(Subramanian & Bhat 1989, as S. malabarica); B. S. goanensis (Bhat & Kendrick 1993);
C. S. guangxiensis (Xia et al. 2014); D. S. malloti (Xia et al. 2015); E. S. lushanensis (this work);
E S. pluriseptata (Monteiro et al. 2016). Scale bars = 20 um.
Fig. 2. Sporidesmiopsis spp., representative conidia (re-drawn from the literature). A. S. dennisii
Sporidesmiopsis lushanensis sp. nov. (China) ... 879
Key to species of Sporidesmiopsis
IG oinidiaee EO eUSepialen Shia etta : SE Ey AB ete ook heey hte ass 2
Ie Gromidiiass 7eeuseptate f. © eyo eitaae yrs ot eis he cis « Pea ee te eg oe ny ee a 3
2. Conidia 69-92 x 9-13 um, 10-15-euseptate ............... 0 eee eee S. dennisii
2. Conidia 80-130 x 6-8.5 um, 10-15-euseptate ................... S. pluriseptata
3. Conidia fusiform, 30-50 x 11.5-15.5 um, 6-7-euseptate........... S. lushanensis
3/@onidia- obclavate todusitorin, <1 5 ani-digmiom ieee ned ee eee Cetdameres |
4. Conidia 20-30 x 5-7 um, 3-4-euseptate .............. 0. ee eee eee S. goanensis
4S Conidia 2S 2M IONe 13% Liat tion Phase adhe sae Es eee Pees Bho Stites 5
5. Conidia 32-40 x 6-7.5 um, 5-6-euseptate ........ ee eee eee eee S. guangxiensis
5. Conidia 35-60 x 8.5-11.5 um, 3-7-euseptate ......... eee eee ee eee S. malloti
Acknowledgments
The authors express gratitude to Dr. Jian Ma and Dr. Rafael F. Castafteda-Ruiz for
serving as pre-submission reviewers and for their valuable comments and suggestions.
This project was supported by the National Natural Science Foundation of China
(Nos. 30600003, 31093440, 31230001) and the Ministry of Science and Technology of
the People’s Republic of China (Nos. 2006FY120100).
Literature cited
Bhat DJ, Kendrick B. 1993. Twenty-five new conidial fungi from the Western Ghats and the
Andaman Islands (India). Mycotaxon 49: 19-90.
Monteiro JS, Gusmao LFP, Castafeda-Ruiz RF. 2016. Pleurothecium bicoloratum & Sporidesmiopsis
pluriseptata spp. nov. from Brazil. Mycotaxon 131: 145-152. https://doi.org/10.5248/131.145
Santa Izabel TS, Cruz ACR, Gusmao LFP. 2013. Conidial fungi from the semi-arid Caatinga
biome of Brazil. Ellisembiopsis gen. nov., new variety of Sporidesmiella and some notes on
Sporidesmium complex. Mycosphere 4: 156-163. https://doi.org/10.5943/mycosphere/4/2/1
Subramanian CV, Bhat DJ. 1989 [“1987”]. Hyphomycetes from South India I. Some new taxa.
Kavaka 15: 41-74.
Wongsawas M, Wang HK, Hyde KD, Lin FC. 2008. New and rare lignicolous hyphomycetes
from Zhejiang Province, China. Journal of Zhejiang University ScIENcE B - Biomedicine &
Biotechnology 9: 797-801. https://doi.org/10.1631/jzus.B0860008
Xia JW, Ma LG, Castafeda-Ruiz RF, Zhang XG. 2014. A new species of Sporidesmiopsis and three
new records of other dematiaceous hyphomycetes from southern China. Nova Hedwigia 98:
103-111. https://doi.org/10.1127/0029-5035/2013/0145
Xia JW, Ma YR, Gao JM, Li Z, Zhang XG. 2015. Sporidesmiopsis malloti sp. nov. and new records
from southern China. Mycotaxon 130: 827-833. https://doi.org/10.5248/130.827
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 881-884
https://doi.org/10.5248/132.881
Repetophragma elegans sp. nov.
from Hainan Province, China
XIAO-MEI WANG’, SHAN-SHAN CHEN’, XIAO-MAN LIU’, ZI-JIAN ZHAO’,
Hao-Hua Lr, X1u-Guo ZHANG? * & JI- WEN XIA?®
‘College of Agronomy, Jilin Agricultural University, Changchun, Jilin, 130118, China
? Shandong Provincial Key Laboratory for Biology of Vegetable Diseases and Insect Pests,
College of Plant Protection, Shandong Agricultural University,
Taian, Shandong, 271018, China
* CORRESPONDENCE TO: “sdau613@163.com, * xiajiwen1@126.com
ABSTRACT—A new species, Repetophragma elegans, is described and illustrated from
specimens collected on dead branches in Hainan Province, China. The fungus is characterized
by subcylindrical to subfusiform 14—18-euseptate brown conidia with rounded apices, apical
mucilaginous tunica, and truncate bases.
Key worps—conidial fungi, taxonomy
Introduction
During our ongoing survey of microfungi associated with woody debris
in tropical forests of Hainan Province, China, an undescribed species with
the morphological characteristics of the genus Repetophragma Subram. was
collected on the dead stems of an unidentified broadleaf tree.
Materials & methods
Collected samples of woody debris were placed in separate zip-lock plastic bags,
taken to the laboratory, and incubated at 27°C for more than 2 weeks in an artificial
climate box in 9-cm-diam. plastic Petri dishes containing moistened filter paper. Samples
were examined under an Olympus SZ61 dissecting microscope. At least 50 mature
conidia and 30 conidiophores were mounted in lactophenol, measured at 600x and
1000x magnifications, and photographed with an Olympus BX53 microscope. Adobe
882 ... Wang & al.
Fic. 1. Repetophragma elegans (holotype, HSAUP H6478).
Conidiophores, conidiogenous cells, and conidia.
Repetophragma elegans sp. nov. (China) ... 883
Photoshop 7.0 was used to prepare the photographic images, and the backgrounds
were replaced for esthetic reasons. Type specimens were deposited in the Herbarium
of Department of Plant Pathology, Shandong Agricultural University, Taian, China
(HSAUP) and the Mycological Herbarium, Institute of Microbiology, Chinese Academy
of Sciences, Beijing, China (HMAS).
Taxonomy
Repetophragma elegans J.W. Xia & X.G. Zhang, sp. nov. FIG. 1
MycoBAank MB 821043
Differs from Repetophragma calongei by its wider conidia, with an apical mucilaginous
tunica.
Type: China, Hainan Province, Diaoluo Mountain, on dead stems of an unidentified
broadleaf tree, 10 Apr. 2014, J.W. Xia (Holotype, HSAUP H6478; isotype, HMAS
245642).
EryMoLoey: elegans, referring to elegant, describing the conidia.
COLONIES on natural substrate effuse, brown to dark brown, hairy. Mycelium
partly superficial, partly immersed in the substrate, composed of septate, pale
brown, smooth, 2-4 um diam. hyphae. CoNIDIOPHORES macronematous,
mononematous, unbranched, erect, straight to slightly flexuous, cylindrical,
smooth, thick-walled, brown to dark brown, up to 5-septate, 13.5-50 x 6.5-9.5
tm. CONIDIOGENOUS CELLS monoblastic, terminal, indeterminate, brown to
dark brown, cylindrical, with 1-5 annellidic percurrent extensions, 6-13.5 x
6-7 um. Conidial secession schizolytic. Conrp1a solitary, straight or slightly
curved, subcylindrical to subfusiform, smooth-walled, brown to dark brown,
14-18-euseptate, 75-115 x 12.5-15.5 um (excluding mucilage), apex rounded,
with an apical mucilaginous tunica, truncate at the base, 6-7 um diam.
COMMENTS—Repetophragma was introduced by Subramanian (1992) to
accommodate Sporidesmium-like species that produced conidiogenous cells
with annellidic percurrent extensions and euseptate conidia. Castaheda-Ruiz
et al. (2011) redisposed taxa in Repetophragma. According to Index Fungorum
(2017) a total of 35 species are included in Repetophragma.
Repetophragma elegans is morphologically similar to R. calongei J. Mena
et al. in conidial shape. However, R. calongei can be easily separated by its
narrower conidia (8-10.5 um) without an apical mucilaginous tunica (Silvera-
Simon et al. 2009).
Acknowledgments
The authors express gratitude to Dr. Rafael E Castafieda-Ruiz and Dr. Jian Ma for
serving as pre-submission reviewers and for their valuable comments and suggestions.
884 ... Wang & al.
This project was supported by the National Natural Science Foundation of China (Nos.
30600003, 31093440, 31230001, 31493010, 31493011) and the Ministry of Science and
Technology of the People’s Republic of China (No. 2006FY 120100).
Literature cited
Castafieda-Ruiz RF, Heredia G, Arias RM, McKenzie EHC, Hyde KD, Stadler M, Saikawa M,
Gené J, Guarro J, Iturriaga T, Minter DW, Crous PW. 2011. A new species and re-disposed taxa
in Repetophragma. Mycosphere 2: 273-289.
Index Fungorum. 2017. http://indexfungorum.org/names/names.asp (Accessed: 30 May 2017).
Silvera-Sim6n C, Mena-Portales J, Gené J, Cano J, Guarro J. 2009. Repetophragma calongeii sp. nov.
and other interesting dematiaceous hyphomycetes from the North of Spain. Anales del Jardin
Botanico de Madrid 66: 33-39. https://doi.org/10.3989/ajbm.2218
Subramanian CV. 1992. A reassessment of Sporidesmium (hyphomycetes) and some related taxa.
Proceeding of the Indian National Science Academy, B 58: 179-190.
MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 885-893
https://doi.org/10.5248/132.885
A contribution to the study of Helotiales and
Rhytismatales in Turkey
MAKBULE ERDOGDU',, GOKHAN DOGAN’,
ELSAD HUSEYIN' & ZEKIYE SULUDERE”
' Ahi Evran University, Faculty of Science and Literature, Department of Biology,
Bagbasi, Kirsehir, Turkey
?Gazi University, Faculty of Science, Department of Biology,
Teknikokullar, Ankara, Turkey
* CORRESPONDENCE TO: merdogdu@ahievran.edu.tr
ABSTRACT—Naemacyclus fimbriatus, Lophodermium juniperinum, and Marssonina daphnes
have recently been discovered in Turkey. This is the first record of Naemacyclus from Turkey.
Morphological data obtained by light and scanning electron microscopy of these fungi are
presented.
Key worps—acervular anamorph, Ascomycota, new host, new records, SEM
Introduction
Rhytismatales are an order of endophytic, parasitic, or saprotrophic fungi in
Leotiomycetes (Ascomycota), the inoperculate discomycetes. Especially common
on conifers, grasses and members of Ericaceae, they are also found on other
vascular plants. The species disperse by ascospores and at least in temperate
regions usually infect their hosts in spring/summer to develop fruiting bodies
the next year on dead material (Lantz et al. 2011). The order includes plant
parasitic fungi causing serious needle cast, such as Lophodermium seditiosum
Minter et al. on Pinus sylvestris (Minter 1981b)., Rhytismatalean fungi the
members of Rhytismatales are poorly known in Turkey and have not been yet
intensively studied.
Within Leotiomycetes, Helotiales represents the largest order of inoperculate
discomycetes—an ecologically and morphologically highly diverse group of
886 ... Erdogdu & al.
ascomycetes that also includes lichen-inhabiting (lichenicolous) species (Suija
et al. 2015). At present the order comprises c. 4000 species of saprophytes,
mycorrhizal fungi, root endophytes, and plant and fungal (including lichens)
parasites (Schoch et al. 2009).
Materials & methods
Plant specimens infected with microfungi were collected from Erciyes Mountain in
Kayseri province of Turkey. The host specimens were prepared following conventional
herbarium techniques. Host plants were identified using the FLORA OF TURKEY AND
East AEGEAN IsLANDS (Davis 1965-85). Thin fungal sections prepared from host
tissue were examined under a Leica DM E light microscope and measured from mounts
in 5% KOH or tap water. Close-up photographs of infected host surface were done via
Leica EZ4D stereomicroscope. The fungi were identified using relevant literature (for
Naemacyclus — DiCosmo et al. 1984, Hou et al. 2006; for Lophodermium -— Dennis
1981, Ellis & Ellis 1987; for Marssonina - Grove 1937, Ellis & Ellis 1987, Ignatavicitité &
Treigiené 1998). All examined specimens were deposited in the Mycology Laboratory
of Ahi Evran University, Department of Biology, Kirsehir, Turkey (C), under Gdkhan
Dogan (GD) collection numbers.
For scanning electron microscopy (SEM), 8-10 mm square pieces of infected leaf
or cone scale were mounted on aluminium stubs with double-sided adhesive tape,
coated with gold using Polaron SC 502 Sputter Coater, and examined using a Jeol JSM
6060 scanning electron microscope operated at 5-10 kV in the Electron Microscopy
Unit, Gazi University (Turkey).
Taxonomy
Naemacyclus fimbriatus (Schwein.) DiCosmo, Peredo & Minter,
Eur. J. For. Path. 13(4): 207 (1983) Pr
APOTHECIA scattered, immersed to erumpent, sessile, circular to
subcircular, 300-430 x 230-285 um, dark brown to black. Asci cylindrical,
short-stalked, thin-walled, 85-95 x 8.5-9.5 um, rostrate at the apex, without
circumapical thickening, discharging spores through a small apical pore,
8-spored. Ascospores fasciculate, filiform, (50-)54-80 x 2-2.2 um, rounded
at both ends, hyaline, (2—)5-septate, rarely aseptate, with mucous sheath.
PaRAPHYSES filiform, unbranched, septate, 90-105 x 1 um, hyaline, covered by
a thin mucous sheath.
SPECIMEN EXAMINED—TURKEY, KayseErI, Erciyes mountain, Turkish World Forest,
38°36'12”N 35°3056”E, 1850-1900 m asl., on fallen female cone scales of Pinus nigra L.
(Pinaceae), 31.05.2010, G. Dogan (AEUT GD 1069).
Notes: While DiCosmo et al. (1983) resolved the nomenclatural problems
surrounding N. fimbriatus, the systematic position of this species is still
Naemacyclus, Lophodermium & Marssonina spp. new for Turkey ... 887
Tay Fi i
AN PY
Dae i
PLaTE 1. Microscopic characters of Naemacyclus fimbriatus. A: apothecia on cone scales;
E: asci and
vertical section of an apothecium;
D:
G: ascospore;
>
C: apothecia on cone scales (SEM);
2
B
H: lacto phenol cotton blue mount
F-H: 15 um.
F: ascus and ascospores;
showing ascospore. Scale bars = D: 100 um;
ascospores;
E20 pm;
888 ... Erdogdu & al.
controversial. Naemacyclus fimbriatus was once placed in the Phacidiaceae,
but DiCosmo (1979) suggested that it is more closely related to Rhytismataceae
because of J- apical ring and ascospores covered by a mucous sheath (Kirk
et al. 2001, Cannon & Kirk 2007). The recent analysis of the partial small
subunit rDNA (Hou et al. 2006) indicates that N. fimbriatus is closely related
to taxa in the Helotiales.
Although our Turkish specimen of Naemacyclus fimbriatus is
morphologically similar to specimens described in literature (DiCosmo et
al. 1984, Vujanovic et al. 1998, Hou et al. 2006), it does differ slightly in some
aspects, particularly with respect to ascospore size and number of septa:
75-90 x 2-2.5 um and 7-septate (DiCosmo et al. 1984); 78-91 x 2-2.5 um
and 7-septate (Vujanovic et al. 1998); 65-95 x 1-1.5 um and (2—)4—6-septate
(rarely aseptate) (Hou et al. 2006). The ascospores in the specimen from
Turkey are shorter and (2—)5-septate (rarely aseptate).
Naemacyclus fimbriatus is widely distributed on Pinus spp. in Asia (China),
Europe, and North America (Cannon et al. 1985; Dennis 1981; Dudka et al.
2004; Eriksson 2014; Gremmen 1960, as Lasiostictis fimbriata; Hanlin 1963,
as Stictis fimbriata; Lin 2012; Minter 1981a; Sherwood 1979, as Lasiostictis
fimbriata; Vujanovic et al. 1998). The genus Naemacyclus and N. fimbriatus
are reported for the first time from Turkey.
Lophodermium juniperinum (Fr.) De Not., G. Bot. Ital. 2(7-8): 46 (1847) PL. 2
APOTHECIA hysterioid, elliptical, blister-like, strongly raised above the
surface of needles, 600-650 x 200-220 um, blackish; disc soft, whitish. Asc1
cylindrical-clavate, short-stalked, thin-walled, 95-110 x 13.5-15.5 um,
8-spored; Ascospores fasciculate, filiform, attenuated at both ends, 65-87 x
1.5-3 um, guttulate, hyaline, with a thin hyaline gelatinous coating. PARAPHYSES
slender, filiform, hyaline, swollen and curled at the tip.
SPECIMEN EXAMINED—TURKEY, KaysErI, Erciyes mountain, Develi, 2000-2050 m
asl., conifer plantation area, on dead leaves of Juniperus communis L. (Cupressaceae),
25.07.2011, G. Dogan (AEUT GD1106).
Notes: Ten rhytismataceous ascomycetes from the genera Coccomyces,
Colpoma, Hypoderma, Lophodermium, Pseudophacidium, Soleella, and
Tryblidiopsis have been reported on species of Juniperus; those on juniper
needles are mostly pathogenic, causing needle cast (Hou et al. 2005).
Lophodermium juniperinum is probably weakly parasitic, capable of endophytic
growth in needles for part of its life cycle, like most species of Lophodermium
(Minter 1981b).
Naemacyclus, Lophodermium & Marssonina spp. new for Turkey ... 889
igauy
were
Ss. st
MIOD 1d0e= qurcer
PLATE 2. Microscopic characters of Lophodermium juniperinum. A: apothecia on leaf;
B: apothecium on leaf (SEM); C-E: vertical section of an apothecium (SEM); F: vertical section
of an apothecium; G: ascus and ascospores; H: ascospores. Scale bars = F: 200 um; G,H: 25 um.
890 ... Erdogdu & al.
The Turkish specimen of L. juniperinum agrees with other reports of the
species in ascomata, asci and ascospores morphology; the only observable
difference being the smaller dimensions of ascomata, asci, and ascospores.
Dennis (1981) describes 1 x 0.4 mm apothecia, 130 x 17 um asci, and
60-100 x 2 um ascospores; Ellis & Ellis (1987) cite 1 mm long apothecia
and 60-90 x 2 um ascospores.
Lophodermium juniperinum is known from Asia (Dudka et al. 2004),
Europe, and North America (Hou et al. 2005). This species is reported for
the first time from Turkey.
Marssonina daphnes (Roberge ex Desm.) Magnus, Hedwigia 45: 89 (1906) PL. 3
LEAF SPOTS amphigenous, irregularly rounded, 1-3 mm diam.,
greenish and then brownish. Conrpiomata acervular, amphigenous, at
first immersed in host tissue, later erumpent, small, pale brown. CoNIDIA
ovoid to pyriform, slightly curved, 15-20 x 6.5-8 um, when mature with
transverse septum near the base, not constricted, obtuse at the apex,
truncate at the base, with numerous guttulations, hyaline.
SPECIMEN EXAMINED—TURKEY, KaysERI, Erciyes mountain, Kayseri memorial
forest, 38°36'13”N 35°30’58’”E, 1870 m asl., on living leaves of Daphne oleoides
Schreb. subsp. oleoides (Thymelaeaceae), 25.07.2011, G. Dogan (AEUT GD1080).
Notes: Marssonina daphnes is the causal agent of the daphne anthracnose,
which usually causes little damage. The Turkish specimen of Marssonina
daphnes is morphologically similar to specimens described in literature
(Grove 1937, Ellis & Ellis 1987, Ignatavicitite & Treigiené 1998). However,
the conidia of the Turkish samples are wider than the British collection
(12-20 x 4-5 um; Grove 1937), while those reported by Ignataviciiité &
Treigiené (1998) are slightly narrower and shorter than our specimen.
Marssonina daphnes is distributed on Daphne spp. in Asia (Japan),
Australasia, Europe, and North America (Conners 1967; Ginns 1986; Cook
& Dubé 1989; Kobayashi 2007; Pennycook & Galloway 2004; Piatek &
Wolczanska 2004; Shaw 1973). The species is reported for the first time
from Turkey, and Daphne oleoides Schreb. subsp. oleoides represents a new
host record.
Acknowledgments
We are grateful to Dr. Eugene Yurchenko, and Sevda Kirbag for serving as pre-
submission reviewers, to Dr. Pennycook for nomenclatural review.
Naemacyclus, Lophodermium & Marssonina spp. new for Turkey ... 891
<
~
ao ocd ers
. —-<———<
PLATE 3. Microscopic characters of Marssonina daphnes. A,B: acervulus on leaf (SEM);
C-E: Conidia (SEM); F: Conidia. Scale bar = 10 um.
Literature cited
Cannon PF, Kirk PM. 2007. Fungal families of the world. CAB International Wallingford.
https://doi.org/10.1079/9780851998275.0000
Cannon PF, Hawksworth DL, Sherwood-Pike MA. 1985. The British Ascomycotina. An
annotated checklist. Commonwealth Mycological Institute, Kew.
Conners IL. 1967. An annotated index of plant diseases in Canada and fungi recorded on plants
in Alaska, Canada and Greenland. Canada Department of Agriculture, Ottawa.
Cook RP, Dubé AJ. 1989. Host-pathogen index of plant diseases in South Australia. Field Crops
Pathology Group, South Australian Department of Agriculture, Adelaide.
892 ... Erdogdu & al.
Davis PH (ed.). 1965-85. Flora of Turkey and East Aegean Islands. Vols 1-9. Edinburgh
University Press, Edinburgh.
Dennis RWG. 1981. British ascomycetes. J. Cramer, Vaduz.
DiCosmo F. 1979. Lasiostictis reassessed. Canadian Journal of Botany 57: 1838-1840.
https://doi.org/10.1139/b79-229
DiCosmo F, Peredo H, Minter DW. 1983. Cyclaneusma gen. nov., Naemacyclus and Lasiostictis,
a nomenclatural problem solved. European Journal of Forest Pathology 13(4): 206-212.
https://doi.org/10.1111/j.1439-0329.1983.tb00119.x
DiCosmo F, Nag Raj TR, Kendrick WB. 1984. A revision of the Phacidiaceae and related
anamorphs. Mycotaxon 21: 1-234.
Dudka IO, Heluta VP, Tykhonenko YY, Andrianova TV, Hayova VP, Prydiuk MP, Dzhagan VV,
Isikov VP. 2004. Fungi of the Crimean Peninsula. M.G. Kholodny Institute of Botany, Ukraine.
Ellis MB, Ellis JP. 1987. Microfungi on land plants: an identification handbook. Croom Helm,
London-Sydney.
Eriksson OE. 2014. Checklist of the non-lichenized ascomycetes of Sweden. Symbolae Botanicae
Upsalienses 36(2). 501 p.
Ginns JH. 1986. Compendium of plant disease and decay fungi in Canada 1960-1980. Canadian
Government Publishing Centre, Ottawa. https://doi.org/10.5962/bh1.title.58888
Gremmen J. 1960 [“1959”]. A contribution to the mycoflora of pine forests in the Netherlands.
Nova Hedwigia 1(3-4): 251-288.
Grove WB. 1937. British stem- and leaf-fungi (Coelomycetes), vol. 1. Sphaeropsidales. Cambridge
University Press, Cambridge.
Hanlin RT. 1963. A revision of the ascomycetes of Georgia. Georgia Agricultural Experiment
Station, Mimeo Series n.s. 175. 65 p.
Hou CL, Gao J, Piepenbring M. 2006. Four rhytismataceous ascomycetes on needles of pine from
China. Nova Hedwigia 83: 511-522. https://doi.org/10.1127/0029-5035/2006/0083-0511
Hou CL, Lin YR, Piepenbring M. 2005. Species of Rhytismataceae on needles of Juniperus spp.
from China. Canadian Journal of Botany 83: 37-46. https://doi.org/10.1139/b04-149
Ignataviciiiteé M, Treigiené A. 1998. Mycota Lithuaniae, vol. 9. Melanconiales. UAB Vaslstieciu
Laikrastis, Vilnius.
Lantz H, Johnston PR, Park D, Minter DW. 2011. Molecular phylogeny reveals a core clade of
Rhytismatales. Mycologia 103: 57-74. https://doi.org/10.3852/10-060
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s dictionary of the fungi.
9th edn. CAB International, Kew.
Kobayashi T. 2007. Index of fungi inhabiting woody plants in Japan. Host, distribution and
literature. Zenkoku-Noson-Kyoiku Kyokai Publishing Co., Tokyo.
Lin YR. 2012. Rhytismatales. Flora Fungorum Sinicorum 40. 261 p.
Minter DW. 1981a. Microfungi on needles, twigs and cones of pines in Czechoslovakia. Ceska
Mykologie 35(2): 90-101.
Minter DW. 1981b. Lophodermium on pines. Mycological Papers 147. 54 p.
Piatek M, Wolczanska A. 2004. Some phytopathogenic fungi rare or new to Poland. Polish
Botanical Studies 49(1): 67-72.
Pennycook SR, Galloway DJ. 2004. Checklist of New Zealand “fungi” 401-488, in: EHC
McKenzie (ed.). Introduction to fungi of New Zealand. Fungi of New Zealand vol. 1. Fungal
Diversity Research Series 14.
Schoch CL, Sung GH, Lépez-Giraldez FE, Townsend JP, Miadlikowska J. Hofstetter V,
Robbertse B, Matheny PB, Kauff F, Wang Z, Gueidan C, Andrie RM, Trippe K, Ciufetti
Naemacyclus, Lophodermium & Marssonina spp. new for Turkey ... 893
LM, Wynns A, Fraker E, Hodkinson BP, Bonito G, Groenewald JZ, Arzanlou M, de Hoog
GS, Crous PW, Hewitt D, Pfister DH, Peterson K, Gryzenhout M, Wingfield MJ, Aptroot
A, Suh SO, Blackwell M, Hillis DM, Griffith GW, Castlebury LA, Rossman AY, Lumbsch
HT, Licking R, Biidel B, Rauhut A, Diederich P, Ertz D, Geiser DM, Hosaka K, Inderbitzin
P, Kohlmeyer J, Volkmann-Kohlmeyer B, Mostert L, O’Donnell K, Sipman H, Rogers JD,
Shoemaker RA, Sugiyama J, Summerbell RC, Untereiner W, Johnston PR, Stenroos S, Zuccaro
A, Dyer PS, Crittenden PD, Cole MS, Hansen K, Trappe JM, Yahr R, Lutzoni F, Spatafora
JW. 2009. The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and
evolution of fundamental reproductive and ecological traits. Systematic Biology 58: 224-239.
https://doi.org/10.1093/sysbio/syp020
Shaw CG. 1973. Host fungus index for the Pacific Northwest - I. Hosts. Washington
Agricultural Experimental Station Bulletin 765. 121 p.
Sherwood MA. 1979. Phacidiales Exsiccati. Decades I-III. Mycotaxon 10: 241-245.
Suija A, Ertz D, Lawrey JD, Diederich P. 2015. Multiple origin of the lichenicolous life
habit in Helotiales, based on nuclear ribosomal sequences. Fungal Diversity 70: 55-72.
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https://doi.org/10.1094/PDIS.1998.82.8.959A
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 895-901
https://doi.org/10.5248/132.895
Huneckia pollinii and Flavoplaca oasis
newly recorded from China
CoNnG-CoNnG MIA0™, XIANG-XIANG ZHAO”,
ZUN-TIAN ZHAO’, HURNISA SHAHIDIN? & Lu-LU ZHANG"
' Key Laboratory of Plant Stress Research, College of Life Sciences,
Shandong Normal University, Jinan, 250014, P. R. China
? Lichens Research Center in Arid Zones of Northwestern China,
College of Life Science and Technology, Xinjiang University,
Xinjiang , 830046 , PR. China
“ CORRESPONDENCE TO: ccmjy123@163.com
AsstRact—Huneckia pollinii and Flavoplaca oasis are described and illustrated from
Chinese specimens. The two species and the genus Huneckia are recorded for the first time
from China.
Keyworps—Asia, lichens, taxonomy, Teloschistaceae
Introduction
Teloschistaceae Zahlbr. is one of the larger families of lichenized fungi. It
includes three subfamilies, Caloplacoideae, Teloschistoideae, and Xanthorioideae
(Gaya et al. 2012; Arup et al. 2013). Many new genera have been proposed based
on molecular phylogenetic investigations (Arup et al. 2013; Fedorenko et al.
2012; Gaya et al. 2012; Kondratyuk et al. 2013, 2014a,b, 2015a,b,c,d). Currently,
the family contains approximately 79 genera (Karnefelt 1989; Arup et al. 2013;
Kondratyuk et al. 2013, 2014a,b, 2015a,b,c,d; Sochting et al. 2014a,b).
Huneckia S.Y. Kondr. et al. was described in 2014 (Kondratyuk et al. 2014a)
based on morphological, anatomical, chemical, and molecular data. It is
characterized by continuous to areolate thalli, paraplectenchymatous cortical
* CONG-CONG MIAO & XIANG-XIANG ZHAO contributed equally to this research.
896 ... Miao, Zhao & al.
layers, biatorine apothecia, Blastenia-type true exciples, narrowly bacilliform
conidia, and the presence of chrysophanol, rhein, and chrysophanal. Huneckia
is similar to Blastenia A. Massal. but differs by having ascospores with very
thick cell walls at the poles and in the secondary chemistry (Kondratyuk et
al. 2014a). Huneckia includes two species, H. pollinii and H. rheinigera (Elix &
S.Y. Kondr.) S.Y. Kondr et al. (Kondratyuk et al. 2014a). No species of Huneckia
have been reported from China .
Flavoplaca Arup etal., described in 2013 (Arup etal. 2013), is characterized by
yellow crustose or squamulose thalli, orange zeorine apothecia, polaribilocular
spores with medium to long spore septa, and bacilliform to ellipsoid conidia.
The genus is rather similar to Xanthoria (Fr.) Th. Fr. and Caloplaca Th. Fr. in
some characters, but forms a very well delimited clade with many species that
are often sorediate. Flavoplaca includes about 27 species (Arup et al. 2013).
During our research on the taxa of Teloschistaceae in China, we identified
two species as new to the country: Huneckia pollinii and Flavoplaca oasis.
Materials & methods
The specimens, which were collected in Guizhou Province and Inner Mongolia,
are preserved in the Lichen Section of the Botanical Herbarium, Shandong Normal
University, Jinan, China (SDNU) and the Lichens Research Center in Arid Zones of
Northwestern China, Xinjiang University, Wulumugi, China (XJU). The morphological
and anatomical characters were examined under a COIC XTL7045B2 stereo microscope
and an Olympus CX41 polarizing microscope. Thalli and medullae were tested with K
(10% aqueous solution of potassium hydroxide) and C (a saturated solution of aqueous
sodium hypochlorite) for identification. Calcareous and non-calcareous rocks were
determined by application of concentrated HCl. The lichen substances were identified
using standardized thin layer chromatography techniques (TLC) with solvent system C
(Orange et al. 2010). Photos were taken using Olympus SZX16 stereo and BX61 research
microscopes attached to an Olympus DP72 camera.
Taxonomic descriptions
Huneckia pollinii (A. Massal.) S.Y. Kondr., Karnefelt, A. Thell, Elix, Jung Kim,
A.S. Kondr. & Hur, Acta Bot. Hung. 56: 111 (2014). Pie. tT
MorPHOLOGY—THALLUS crustose, continuous, very thin to areolate at
margin, grey to whitish grey. APOTHECIA biatorine, sessile, 0.3-1.0 mm diam.;
disc dark brown to rust-brown; proper exciple consisting of radiating, thick
walled long-septate cells, prosoplectenchymatous; exciple and epithecium K+
reddish violet; without crystals; epihymenium brown; hymenium colourless
to pale brown, 74-90 um high; hypothecium sand yellow, roundish to
irregular shape, without oil droplets; paraphyses 2.2-3.0 um wide, upper part
Huneckia & Flavoplaca lichens new for China ... 897
Fic. 1. Huneckia pollinii (SDNU 20160195). A: thallus; B: apothecia; C: apothecium section;
D: strong red turning to purple K reaction of epihymenium; E: ascospores; F: conidia.
pigmented, without swollen tip, abundantly branching at tips. Asci 8-spored,
Teloschistes-type; ascospores polaribilocular, ellipsoid, mostly “sand-clock”-
shaped; 13.2-19 x 7.5-9.6 um, with cell walls 1-2(-2.5) um thick (thickest
at the poles); spore septum 2.9-5.7 um. Conip1A elliptical to narrowly
bacilliform, 1.5-2.2 x 3.8-5.1 um.
CuHEMISTRY— thallus and medulla K-, C-. Apothecia K+ purple, C+ red.
Epihymenium K+ strong red, turning purple with coloured solution. Unknown
substances 1 & 2 (TLC solvent system C Rf. 48 and Rf. 53).
898 ... Miao, Zhao &al.
SUBSTRATE—On wood or on smooth bark.
DistRIBUTION—Huneckia pollinii has been reported from the eastern
United States, Europe, and Mexico (Wetmore 1994, Kondratyuk et al. 2014a).
New to China.
SPECIMENS EXAMINED: CHINA. GUIZHOU, Dejiang, Quankou town alt. 873 m, on bark,
2 June 2016, X.X. Zhao 20160195, 20160180 (SDNU).
ComMMENTS—Caloplaca pollinii var. major B. de Lesd. is similar to Huneckia
pollinii, but differs in having concave brown young apothecia, black immarginate
older apothecia, and larger spores (Bouly de Lesdain 1949). Specimens of
Caloplaca asserigena (J. Lahm) Della Torre & Sarnth. that may be confused
with Huneckia pollinii, can be distinguished by rust-red to red-brown discs and
smaller ascospores.
Flavoplaca oasis (A. Massal.) Arup, Frédén & Sochting,
Nordic J. Bot. 31: 46 (2013). ewe
MorPHOLOGY—THALLUS usually visible but thin, with only some small
granules or areole visible at thallus margin, pale yellow; alveolate cortex; algal
layer always separated by fungal hyphae; without prothallus and vegetative
propagules; isidia, soredia, and blastidia absent; medulla white. APOTHECIA
pseudolecanorine or zeorine type, abundant, sessile, round or elliptical,
0.1-0.5 mm diam.; disc slightly convex, orange to dark orange, epruinose;
proper margin very thin, slightly raised, paler than disc; amphithecium
+ reduced, with numerous algae, cortex poorly developed; epihymenium
granular, yellow; hymenium hyaline, 75-85 um tall; paraphyses simple
or slightly branched above, 1-2 apical cells thickened, up to 7 um wide;
hypothecium hyaline or inspersed, +roundish, consisting of thin-
walled roundish cells, 53-125 um, without crystals or oil droplets,
prosoplectenchymatous, hyphae irregular, 40-50 um high. ExcrpLe
consisting of thin-walled oval to polygone cells, upper side 38-50 um.
PARAPHYSIS mostly simple, rarely forked, swollen in and near the tip cells,
4.5-5.3 um in wide. Asci 43-52 x 11-21 um, 8-spored, Teloschistes-type.
Ascospores polaribilocular, thin-walled, 8.5-13.5 x (4.0-)4.8-7.6 um,
spore septum wide, >% of spore length, 2.9-4.6 um. PYcNIpIA not observed.
CuHEMISTRY—Thallus K+ purple, medulla K-, epihymenium K+ purple.
Unknown substance (TLC solvent system C Rf. 51).
SUBSTRATE—On pure limestone, concrete.
DISTRIBUTION—Flavoplaca oasis, which is widespread in Europe (Arup
2009), is also reported from western Asia (John et al. 2004) and North Africa
Huneckia & Flavoplaca lichens new for China... 899
SAS
4 . cal
ern
Fic. 2. Flavoplaca oasis (SDNU 20160086). A: thallus; B: apothecia; C: apothecium section; D: ascus
and ascospores; E: paraphyses; F: ascospores.
(Thor & Nascimbene 2010). The species grows mainly on pure limestone,
concrete, and mortar (Arup 2009), where it occurs as a free living or parasitic
lichen growing on endolithic Verrucaria s.lat. (Wilk 2011). New to China.
SPECIMENS EXAMINED: CHINA. GuizHou, Dejiang, Jingangling, alt. 1100 m, on
rocks, 26 May 2016, W.C. Wang , X.X. Zhao 20160086, 20160038, 20160814 (SDNU).
INNER MONGOLIA, Daqing mountain, 41°54.60’N 111°47.71’E. alt. 1970 m, on rocks,
12 August 2014. 20141175-b (XJU).
900 ... Miao, Zhao & al.
ComMMENTS—Flavoplaca oasis is similar to Athallia holocarpa (Hoftm.) Arup et
al., which is distinguished by its more yellow-tinged apothecia with thicker and
more prominent proper margins and spores with a broader isthmus. Specimens
of F. oasis may also be confused with F. polycarpa (A. Massal.) Arup et al.,
which can be distinguished by larger apothecia with thicker margins and more
distinct, thicker orange thalli.
Acknowledgements
We thank Dr. Li-song Wang and Dr. Xin-yu Wang (Key Laboratory for Plant
Diversity and Biogeography of East Asia, Kunming Institute of Botany, CAS)
for providing great help during the study. We also thank Dr. Andre Aptroot
(ABL Herbarium, Soest, Netherlands) and Dr. S.Y. Guo (Key Laboratory of
Systematic Mycology and Lichenology Laboratory, Institute of Microbiology, CAS,
Beijing, China) for presubmission review and providing great guidance during the
study. This work was supported by the National Natural Science Foundation of China
(31570017) and the Scientific Research Foundation of Graduate School of Shandong
Normal University (SCX201629).
Literature cited
Arup U. 2009. The Caloplaca holocarpa group in the Nordic countries, except Iceland.
Lichenologist 41: 111-130. https://doi.org/10.1017/S0024282909008135
Arup U, Sechting U, Frédén P. 2013. A new taxonomy of the family Teloschistaceae. Nordic
Journal of Botany 31: 16-83. https://doi.org/10.1111/j.1756-1051.2013.00062.x
Bouly De Lesdain M. 1949. Notes lichénologiques, N° XX XV. Bulletin de la Société Botanique de
France 96(7-9): 173-175. https://doi.org/10.1080/00378941.1949.10837606
Fedorenko NM, Stenroos S, Thell A, Karnefelt I, Elix JA, Hur JS, Kondratyuk SY. 2012. Molecular
phylogeny of xanthorioid lichens (Teloschistaceae, Ascomycota), with notes on their
morphology. Bibliotheca Lichenologica 108: 45-64.
Gaya E, Hégnabba F, Holguin A, Molnar K, Fernandez-Brime S, Stenroos S, Arup U, Sechting
U, Van den Boom P, Liicking R, Sipman HJM, Lutzoni F. 2012. Implementing a cumulative
supermatrix approach for a comprehensive phylogenetic study of the Teloschistales
(Pezizomycotina, Ascomycota). Molecular Phylogenetics and Evolution 63: 374-387.
https://doi.org/10.1016/j.ympev.2012.01.012
John V, Seaward MRD, Sipman HJM, Zedda L. 2004. Lichens and lichenicolous fungi from Syria,
including a first checklist. Herzogia 17: 157-177.
Karnefelt I. 1989. Morphology and phylogeny in the Teloschistales. Cryptogamic Botany 1:
147-203.
Kondratyuk SY, Lékés L, Tschabanenko S, Haji Moniri M, Farkas E, Wang XY, Oh SO, Hur JS.
2013. New and noteworthy lichen-forming and lichenicolous fungi. Acta Botanica Hungarica
55(3-4): 275-349. https://doi.org/10.1556/ABot.55.2013.3-4.9
Kondratyuk SY, Jeong MH, Yu NN, Karnefelt I, Thell A, Elix JA, Kim JA, Kondratiuk AS, Hur JS.
2014a. A revised taxonomy for the subfamily Caloplacoideae (Teloschistaceae, Ascomycota)
based on molecular phylogeny. Acta Botanica Hungarica 56: 93-123.
https://doi.org/10.1556/ABot.56.2014.1-2.12
Huneckia & Flavoplaca lichens new for China... 901
Kondratyuk SY, Karnefelt I, Thell A, Elix JA, Kim JA, Jeong MH, Yu NN, Hur JS. 2014b. A revised
taxonomy for the subfamily Xanthorioideae (Teloschistaceae, Ascomycota) based on molecular
phylogeny. Acta Botanica Hungarica 56: 141-178.
https://doi.org/10.1556/ABot.56.2014.1-2.12
Kondratyuk SY, Karnefelt I, Thell A, Elix JA, Kim J, Kondratiuk AS, Hur JS. 2015a. Tassiloa,
a new genus in the Teloschistaceae (lichenized Ascomycetes). Graphis Scripta 27(1-2): 22-26.
Kondratyuk SY, L6kés L, Kim JA, Kondratiuk AS, Jeong MH, Jang SH, Oh SO, Hur JS. 2015b.
Three new monotypic genera of the caloplacoid lichens (Teloschistaceae, lichen-forming
Ascomycetes). Mycobiology 43: 195-202. https://doi.org/10.5941/MYCO.2015.43.3.195
Kondratyuk SY, Karnefelt I, Thell A, Elix JA, Kim J, Kondratiuk AS, Hur JS. 2015c. Brownlielloideae,
a new subfamily in the Teloschistaceae (Lecanoromycetes, Ascomycota). Acta Botanica
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Kondratyuk SY, Kim JA, Yu NH, Jeong MH, Jang SH, Kondratiuk AS, Zarei-Darki B, Hur JS.
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https://doi.org/10.15407/ukrbotj72.06.574
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A, Sancho LG, Frédén P, Arup U. 2014b. Charcotiana and Amundsenia, two new genera in
Teloschistaceae (lichenized Ascomycota, subfamily Xanthorioideae) hosting two new species
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species. Lichenologist 46: 763-782. https://doi.org/10.1017/S0024282914000395
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115: 83-98. https://doi.org/10.5248/115.83
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 903-907
https://doi.org/10.5248/132.903
Paliphora bicolorata sp. nov.
from the Brazilian Atlantic Forest
ELAINE MALOSSO",, PHELIPE M.O. Costa’, MARCELA A. BARBOSA’,
GABRIELA V.R. DA SILVA? & RAFAEL F. CASTANEDA-RUIZ?
"Centro de Biociéncias, Departamento de Micologia,
Universidade Federal de Pernambuco, Avenida da Engenharia,
s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil
? Programa de Pés-Graduagao em Biologia de Fungos,
Universidade Federal de Pernambuco, Avenida da Engenharia,
s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil
* Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT),
Alejandro de Humboldt, OSDE, Grupo Agricola,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
* CORRESPONDENCE TO: elaine.malosso@ufpe.br
ABSTRACT—A new species, Paliphora bicolorata, collected from decaying leaves of an
unidentified plant, is described and illustrated. It is characterized by bicolored conidiophores
with 3-4 dark reddish-brown distal cells and golden brown to brown lower cells and
cylindrical-fusiform, 1-septate, hyaline conidia. A key to Paliphora species is provided.
KEY WoRDS—neotropic, asexual fungi, systematics
Introduction
The genus Paliphora Sivan. & B. Sutton was introduced by Sivanesan & Sutton
(1985) with P. aurea Sivan. & B. Sutton as type species; five more species were
added by Kuthubutheen (1987), Alcorn (1996), Gusmao et al. (2008), and Goh
et al. (2014). Paliphora is distinguished by distinct, unbranched, multiseptate,
setiform conidiophores that form hispid golden brown to brown colonies and
polytretic, integrated, determinate, intercalary (rarely terminal) conidiogenous
cells with tretic loci (situated immediately below transverse septa) producing
successive, cylindrical or somewhat subfusiform to subacerose, hyaline,
904 ... Malosso & al.
unicellular or euseptate conidia that accumulate in slimy masses around the
conidiogenous cells. The production of successive conidia in dense mucous
masses or packets through tretic loci is peculiar among hyphomycetes and
serves as the main diagnostic character of Paliphora (Alcorn. 1996, Gusmao et
al. 2008, Sivanesan & Sutton 1985).
During a mycological survey of microfungi associated with leaf litter in a
Brazilian Atlantic forest, we collected a conspicuous fungus, described here as
a new Paliphora species.
Materials & methods
Individual collections were placed in plastic bags, taken to the laboratory, and
treated according to Castafieda-Ruiz et al. (2016). Mounts were prepared in polyvinyl
alcohol-glycerol (8 g PVA in 100 ml of water, plus 5 ml of glycerol) and lactofuchsin
(0.1 g acid fuchsin, 100 ml 85% lactic acid following Carmichael (1955) or in lactic acid
(90%) and measurements were made at a magnification of x1000 under a Nikon Eclipse
Ni-U microscope with bright field optics; photomicrographs were obtained using DIC
optics with a Nikon DS-Fi2 camera. The holotype was deposited in the Herbarium of
Universidade Federal de Pernambuco, Recife, Brazil (URM).
Taxonomy
Paliphora bicolorata Malosso, P.M.O. Costa & R.F. Castafieda, sp. nov. Fie. 1
MycoBAnk MB 823671
Differs from Paliphora intermedia by its bicolored conidiophores.
Type: Brazil, Pernambuco, Igarassu, RPPN Reftigio Ecoldgico Charles Darwin, 7°48’S
34°57’W, on decaying leaves of an unidentified plant, 12.V.2017, coll. RM.O. Costa
(Holotype, URM 90132).
Erymo toey: Latin, bicolorata, two-colored, referred to the conidiophores.
CoLonigs on the natural substrate effuse, hispid, reddish-brown to brown.
Mycelium superficial and immersed, composed of septate, branched, brown,
smooth-walled hyphae, 1.5-2.5 um diam. CONIDIOPHORES macronematous,
setiform, unbranched, erect, straight, cylindrical below, acuminate at the apex,
<17-septate, golden brown below, reddish-dark brown at the four distal cells,
smooth below, verruculose near the apex, 170-220 x 5-6 um, slightly lobed
at the base, 9-13 um wide. CONIDIOGENOUS CELLS polytretic, intercalary,
determinate, incorporated, brown, 7-10 x 4-5 um. Conrp1A solitary,
Fig. 1. Paliphora bicolorata (ex holotype, URM 90132). A. Conidia; B. Conidiogenous cells and
dark reddish-brown distal cell; C. Conidiophore basal cell; D. Intercalary conidiogenous cells
with tretic loci and conidia; E. Conidiophores, conidiogenous cells, and conidia. B-D share the
same scale.
Paliphora bicolorata sp. nov. (Brazil) ... 905
906 ... Malosso & al.
pleurogenous, cylindrical to broad subacerose, attenuate at the base, obtuse at
the apex, straight, 1-euseptate, hyaline, smooth, 18-23 x 2 um, accumulating in
white, slimy masses, more or less near the conidiophores central part.
Notes: Among the six previously described Paliphora species, only
P. intermedia Alcorn and P. porosa Kuthub. are superficially similar to
P. bicolorata by producing 1-septate conidia, but these species are otherwise
clearly distinguished from P. bicolorata. In P. intermedia (which has been
isolated from decaying leaves of several plants in Brazil, Cuba, and Mexico;
Gusmao et al. 2008) the conidia are shorter (14.5-19 um long) with a slightly
asymmetrical septum and the conidiophores are uniformly brown (Alcorn
1996; Rao & de Hoog 1986, as “P. aurea”). Paliphora porosa produces cylindrical
to clavate conidia that are usually 1-septate (rarely 2-3-septate) and somewhat
constricted at the septa and uniformly golden brown longer (200-280 um)
conidiophores with more septa (15-23-septate; Kuthubutheen 1987).
Key to Paliphora species
Nae -Conidianisuallytanicelilaie sh oe ei ek ovat eh or Sear is Sear hs Sea te Sigs sor SIME gr de 2.
Conidiarsuallyseptare 3. Nie Meweg «Meise Whew Micwniaye MEcmiahe Wheelie steriias dst Arwen 3
2. Conidia cylindrical, straight or curved, 17-20 x 1-2 um; conidiophores
cylindrical below, inflated near the apex, verrucose, 13-19-septate,
dark DROW is 142 = 208) 2s UiT ote Sask BES sak Sie Sats cate atin idle cate ego P. inflata
Conidia falcate, obtuse and widest at the apex, attenuating gradually to a narrow
base, 6.5-9 x 1-1.5 um; conidiophores cylindrical, smooth below,
acute, verrucose toward the apex, 11-18(-23)-septate,
evenly golden brown, 99-132 X5.5-6.5 MM ........ eee eee cece eee ee P. aurea
Sa, -OniGiaMhse state: ja uty feycmee fiz catagt a cers Pec coeegl Pe contg) pee heer Pd nee Er orc 4
GOitidta Sel-Sep tate: scare ht gare sahesse, AAT us 28, kT os 28, UOT aya ED be asf ton iat RA eights 6
4. Conidiophores bicolored, golden brown below and dark reddish-brown
at the four distal cells, <17-septate, smooth below, verruculose near the apex,
170-220 x 4-6 um; conidia cylindrical to broad sub-acerose, obtuse
at the apex, attenuated at the base, 18-23 x 2um................. P. bicolorata
Comidiophores:nOt as-abOver eB oe.e Bes ehiee Sad +n Reus Boe oe Sages eee Bad ee Bakes 5
5. Conidiophores evenly brown, cylindrical below, acute at the apex,
16-22-septate, smooth below, verrucose near the apex, 135-205 x 6-7 um;
conidia, cylindrical or fusoid, apex obtuse, base subacute or obconic, straight
or slightly curved, asymmetrically septate, 14.5-19x 2um ....... P. intermedia
Conidiophores evenly golden brown, cylindrical below, acute at the apex,
15-23-septate, smooth below, verrucose near the apex, 200-280 x 5-9 um;
conidia, cylindrical to clavate, symmetrical, slightly constricted at the septa,
apex obtuse, base acute, straight, 12-25 x 1-2.5um ................. P. porosa
Paliphora bicolorata sp. nov. (Brazil) ... 907
6. Conidia 3-6-septate, elongate-fusiform or narrowly, sometimes slightly
sigmoid, 40-70 x 2-3 um; conidiophores cylindrical, apex rounded, brown,
20-63-septate, smooth, 170-540 x4.5-5 um .. 2... eee eee eee eee P. curviapicis
Conidia 8-13-septate, cylindrical, apex obtuse, base obconic, straight or slightly
curved, 50-65 x 2.5 um; conidiophores cylindrical below, bulbous at the basal
cell, apex acute, brown, <16-septate, smooth below, verrucose toward the apex,
BdS 7 NOSG Fas he TE Are altar lt ark elas OM ges P. multiseptata
Acknowledgments
The authors express their sincere gratitude to Dr. Xiu-Guo Zhang and Dr. De-Wei Li
for their critical review of the manuscript. The authors are grateful to Coordena¢ao de
Aperfeicoamento de Pessoal de Nivel Superior (CAPES) for financial support (project
88881.062172/2014-01) and the Programa Ciéncia sem Fronteiras. RFCR is grateful to
the Cuban Ministry of Agriculture and Programa de Salud Animal y Vegetal (project
P131LH003033) for facilities. We acknowledge the facilities provided by Dr. P.M. Kirk
and Drs. V. Robert through the Index Fungorum and MycoBank websites. Dr. Lorelei
Norvell’s editorial review and Dr. Shaun Pennycook’s nomenclature review are greatly
appreciated.
Literature cited
Alcorn JL. 1996. Paliphora intermedia sp. nov. from Australia. Mycotaxon 59: 145-148.
Carmichael JW. 1955. Lacto-fuchsin: a new medium for mounting fungi. Mycologia 47: 611.
Castafieda-Ruiz RF, Heredia G, Gusmao LFP, Li DW. 2016. Fungal diversity of Central and
South America. 197-217, in: DW Li (ed.). Biology of Microfungi. Springer International
Publishing. https://doi.org/10.1007/978-3-319-29137-6_9
Goh TK, Lau WY, Teo KC. 2014. Paliphora curviapicis sp. nov. from Malaysia, and a synopsis of
the genus. Mycotaxon 127: 145-153. https://doi.org/10.5248/127.145
Gusmao LFP, Leao-Ferreira SM, Marques MFO, Almeida DAC. 2008. New species and
records of Paliphora from the Brazilian semi-arid region. Mycologia 100: 306-309.
https://doi.org/10.1080/15572536.2008. 11832485
Kuthubutheen AJ. 1987. Paliphora porosa sp. nov. on leaf litter from Malaysia. Transaction of the
British Mycological Society 89: 270-273. https://doi.org/10.1016/S0007-1536(87)80168-7
Rao V, de Hoog GS. 1986. New or critical hyphomycetes from India. Studies in Mycology 28. 84 p.
Sivanesan A, Sutton BC. 1985. Microfungi on Xanthorrhoea. Transaction of the British
Mycological Society 85: 239-255. https://doi.org/10.1016/S0007-1536(85)80186-8
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 909-918
https://doi.org/10.5248/132.909
Inocybe shawarensis sp. nov. in the Inosperma clade
from Pakistan
A. NASEER’*, A.N. KHALID? & MATTHEW E. SMITH?
‘Centre for Undergraduate Studies & *Department of Botany, University of the Punjab,
Quaid-e-Azam Campus-54590, Lahore, Pakistan
> Department of Plant Pathology, University of Florida, Gainesville, FL, U.S.A.
“CORRESPONDENCE TO: arooj.hons@pu.edu.pk
ABSTRACT —A new species, Inocybe shawarensis, was collected during field research on
ectomycorrhizal fungi associated with oak forests in Swat district, Pakistan. This species,
supported by a combination of morphological and molecular phylogenetic analyses, is
characterized by a brown, fibrillose campanulate pileus, grayish lamellae, slightly bulbous
stipe base, slightly pruinose silvery white stipe, small phaseoliform spores, and clavate
cheilocystidia. We present phylogenetic analyses based on DNA sequences from the internal
transcribed spacer (ITS) region and large subunit (LSU) of the nuclear ribosomal RNA
(rRNA). Phylogenies from both DNA regions cluster I. shawarensis within the Maculata
subclade in the Inosperma clade.
KEY worps — Agaricomycetes, Inocybaceae, Quercus, Shawar Valley
Introduction
The Inocybaceae is one of the most taxonomically diverse families of
Agaricales. Its representatives form ectomycorrhizal (EcM) associations with
many angiosperms and gymnosperms in tropical and temperate areas. Between
70% and 80% of species in the family have been described in association
primarily with ectomycorrhizal plant families Fagaceae, Pinaceae and Salicaceae
(Kirk et al. 2008).
Inocybe (Fr.) Fr., one of three formally described genera in Inocybaceae, was
first established by Fries as a “tribe” of Agaricus in 1821 and later elevated to
genus rank in 1863 (Matheny et al. 2009). The genus has been divided into
910 ... Naseer, Khalid & Smith
subgenera and sections based on spore morphology, shape and distribution of
cystidia on fruit body tissues, and stipe morphology. Multigene phylogenetic
analysis by Matheny (2009) identified seven clades within Inocybaceae: Auritella,
Inocybe, Inosperma, Mallocybe, Mallocybella, Nothocybe, and Pseudosperma.
The Inosperma clade is characterized phylogenetically as a robustly
supported monophyletic group including species with radially fibrillose to
rimose or squamulose pilei and ellipsoid to phaseoliform spores but lacking
metuloid pleurocystidia. Many species of the Inosperma clade also feature
unusual odors, and some may have rubescent or brunnescent flesh.
Among the more than 850 Inocybe species reported worldwide (Matheny et
al. 2009, 2012; Kobayashi & Onishi 2010, Horak et al. 2015, Jabeen et al. 2016),
at least 47 Inocybe species are resolved in the Inosperma clade (Matheny 2009,
Matheny et al. 2009, Kropp et al. 2013, Pradeep et al. 2016, Latha & Manimohan
2016). From Pakistan, 26 species of Inocybe have been reported, but only a few
studies have verified the taxonomic identity of specimens using molecular data
(Ahmad et al. 1997, Ilyas et al. 2013, Saba et al. 2015, Jabeen et al. 2016). The
only species in the Inosperma clade that has been reported from Pakistan is
Inocybe mimica Massee (Saba et al. 2015).
Here we describe a new species, Inocybe shawarensis, from Shawar Valley,
Swat district, Pakistan. The specimens were characterised by morphological
characters as well as molecular datasets based on the internal transcribed spacer
(ITS) and large subunit (LSU) of nrDNA. Based on traditional classifications
(Kithner 1980, Kuyper 1986, Stangl 1989), our new species would be placed in
Inocybe sect. Rimosae within I. subg. Inosperma.
Materials & methods
Sampling Site
The Swat district (34.34-35.55°N 72.08-72.50°E), situated in the Khyber
Pakhtunkhwa Province of Pakistan, is well known for its unique biodiversity (Shinwari
et al. 2003). Its Shawar Valley, which occupies an area of 48.77 km’ within the Hindu
Kush mountain range (Ahmad & Sirajuddin 1996), is topographically mountainous,
varying in elevation from 1200 m to 3800 m (Anonymous 1999). Floristically the Valley
is representative of the western Himalayan Province. The climate is moist temperate
with temperatures averaging a winter minimum of 4.8°C and summer maximum of
33.5°C and an average annual rainfall of about 800 mm, with precipitation occurring in
spring and summer seasons with snowfall at higher elevations (Ullah et al. 2014).
Fruiting bodies of Inocybe, collected during a field investigation of ectomycorrhizal
communities associated with the oaks of Swat during 2014-2016, were found in a pure
Quercus forest in Shawar Valley. The type locality lies in a thick moist temperate forest
Inocybe shawarensis sp. nov. (Pakistan) ... 911
of Quercus oblongata D. Don [= Q. incana Roxb., nom. illeg.] along a tributary of the
Swat River.
Morphological analyses
Fruiting bodies of Inocybe were collected and photographed in the field using a Nikon
D70S camera. Morphological characters were recorded from fresh specimens. Color
designations were based on the Munsell Color System (Munsell 1975). For preservation
specimens were dried using an electric fan heater. Rehydrated material was examined
microscopically in 5% KOH, phloxine, and Melzer’s reagents.
Anatomical features of basidiospores, basidia, cystidia, stipe hyphae and pileus
hyphae were measured at 1000x magnification and include: arithmetic mean of spore
length and width for all spores measured, Q = spore length divided by spore width.
Spore size range was determined by 30 basidiospore measurements from each fruiting
body.
Specimens were deposited in the Herbarium, Department of Botany, University
of the Punjab, Lahore, Pakistan (LAH) and the University of Florida Herbarium,
Gainesville FL, USA (FLAS).
DNA extraction, amplification and sequencing
Genomic DNA was extracted from gill tissue using a modified CTAB method
(Gardes & Bruns 1993). ITS and LSU regions were amplified by the primer pairs ITS1F/
ITS4B and LROR/LRS, respectively. All PCR products were evaluated for successful
amplification using SYBR Green and 1.5% agarose gels with TAE buffer for gel
electrophoresis. Amplicons were prepared for sequencing via enzymatic purification
using Exonuclease I and Shrimp Alkaline Phosphatase enzymes (Werle et al. 1994).
Purified products were sequenced by the University of Florida's Interdisciplinary Center
for Biotechnology Research (http://www.biotech.ufl.edu/). Sequence chromatograms
were trimmed, edited, and assembled using Sequencer 4.1 (GeneCodes, Ann Arbor,
MI). DNA sequences generated for this study were deposited in GenBank.
Molecular phylogenetic analysis
For alignment and phylogenetic analysis, the top 100 BLAST search result sequences
were selected from GenBank using NCBI BLAST (http://www.ncbi.nlm.niih.gov/).
Other closely related species were also included based on the published literature in the
final dataset. Sequences were manually edited and assembled using BioEdit (www.mbio.
ncsu.edu/bioedit/bioedit.html). After sequence alignment by Muscle, all sequences were
trimmed between the conserved motifs 5’-(...GAT) CATTA— and —GACCT(CAAA...)—3’
(Dentinger et al. 2011). Auritella species were selected as outgroup based on results
reported by Larsson et al. (2009). Phylogenetic trees were constructed with the
Maximum Likelihood (ML) algorithm using a general time-reversible model (Nei &
Kumar 2000) and nearest-neighbour interchange as the ML heuristic search method
using MEGA6 software. The topology was assessed by 1000 bootstrap replicates.
Initial tree(s) for the heuristic search were obtained by applying the Neighbor-
Joining method to a matrix of pairwise distances estimated using the Maximum
912 ... Naseer, Khalid & Smith
Composite Likelihood (MCL) approach. The tree is drawn to scale, with branch lengths
measured in the number of substitutions per site. Evolutionary analyses were conducted
in MEGA6 (Tamura et al. 2013).
Taxonomy
Inocybe shawarensis Naseer & Khalid, sp. nov. Fia. 1
MycoBank MB 820130
Differs from Inocybe quietiodor by its brown pileus with a less prominent, campanulate
umbo, its gray gills, its silvery lower stipe and dark gray upper stipe, and its smaller,
more phaseoliform spores.
Type: Pakistan, Khyber Pakhtunkhwa province, Swat district, Shawar Valley, 2100 m
a.s.l, solitary on ground under Quercus oblongata D. Don [= Q. incana Roxb., nom.
illeg.], 14 July 2015, Arooj Naseer ASSW79 (Holotype, FLAS-F-S9456; GenBank
KY616964, KY616966. Isotype, LAH35195; GenBank KY616965).
ErymMo ocy: The specific epithet shawarensis refers to the Shawar Valley, the location
where the type was collected.
Piteus: 30 mm diam., campanulate, umbonate, margin slightly incurved or
deflexed; surface fibrillose, rimose or cracked towards the margin; dark brown
(7.5yR4/8) at the centre, becoming lighter brown in patches towards the margin
that is creamy white (10y8/2). LAMELLAE subdistant, fimbriate, eroded, light
gray (2.5Gy8/2) when young; LAMELLULAE of varying lengths, alternating with
lamellae. STIPE: 4.8 x 0.4 cm, cylindrical, central; surface fibrillose, apex slightly
pruinose, slightly narrower towards apex, base slightly swollen to bulbous; base
light brown (7.5yR4/8) with lower two third creamy or silvery white and upper
one third dark gray (2.5yG5/2)
Basip1ospores [60/2/1], (4.5-)4.7-6.5(-6.8) x (2.7-)2.8 x 3.7(-3.8) um,
avl x avw = 5.2 x 3.2 um, Q = (1.2-)1.3-1.9(-2.5), avQ = 1.63, yellowish
brown in KOH, smooth, thick walled, phaseoliform. BAsrp1a 22-31 x 6-10
um, light brown, clavate, blunt ended, four-spored. CHEILOCYSTIDIA 21-55 x
9-11 um, clavate, thin walled, in groups. PLEUROCYSTIDIA none. PILEIPELLIS
a radially orientated cutis of thin-walled hyphae, 5-7 um diam., cylindrical,
walls smooth or finely encrusted, light brown in KOH, blunt ends, septate,
clamped. CauLocystTip1< only at apex, more or less similar to cheilocystidia.
STIPITIPELLIS hyphae 4-7 um diam., septate, filamentous, unbranched. CLamp
CONNECTIONS present. Odour not recorded.
Molecular phylogenetic analysis
Inocybe shawarensis sequences of the amplified products of ITS (KY616964,
KY616965) and LSU (KY616966) were BLAST searched at NCBI. Both the ITS and
Inocybe shawarensis sp. nov. (Pakistan) ... 913
H o_o
Fic. 1. Inocybe shawarensis (isotype, LAH35195). A, B. basidiomata; C. basidiospores; D. basidia;
E. cheilocystidia; F. caulocystidia; G. pileipellis; H. stipitipellis. Scale bars: A, B = 1.5 um; C = 5 um;
D = 12 um; E, F = 16 um; G, H = 13 um.
914 ... Naseer, Khalid & Smith
LSU sequences suggest a close affinity with I. quietiodor Bon. The LSU sequence
was 97% identical to I. quietiodor (FJ904174) with 100% query coverage, whereas
the ITS sequence showed 93% sequence similarity with I. quietiodor (FJ936168)
with 100% query coverage. The ITS analysis comprised 52 nucleotide sequences
with 1405 positions total in the final dataset, while the LSU analysis comprised 40
nucleotide sequences with 852 positions in the final dataset.
Maculata
Inosperma
JF908117 | geraniodora
EU523576 | mutata
JF908128 | cervicolor _ |Cervicolore:
HQ604816 | calamistrata i
JQ801385 | apiosmota —
AY635766 Auritella aureoplumosa
AY380395 Auritella geoaustralis Jesse
69 AY 380371 Auritella dolichocystis
Fic. 2. Molecular phylogenetic analysis of ITS sequences from Inocybe spp. in the Inosperma
clade inferred by using the Maximum Likelihood method. The analysis involved 52 nucleotide
sequences. The tree with the highest log likelihood (—9223.6738) is shown. Sequences
generated during this study are represented by dots.
Inocybe shawarensis sp. nov. (Pakistan) ... 915
Maculata
EU555463 | cf er Inosperma
JQ994476 | mutata
JN975021 | apiosmota
FN550945 | geraniodora ervicolore
JQ815417 | cervicolor
AY635766 Auritella aureoplumosa
42 AY380395 Auritella geoaustralis
AY635764 Auritella dolichocystis
Out group
0.02
Fic. 3. Molecular phylogenetic analysis of LSU sequences from Inocybe spp. in Inosperma
clade inferred by using the Maximum Likelihood method. The analysis involved 40 nucleotide
sequences. The tree with the highest log likelihood is shown. The sequence generated during
this study is represented by a dot.
Our analysis revealed two major lineages within the Inosperma
clade. The new species clustered with I. quietiodor in the Maculata
subclade, whereas the Cervicolores subclade was resolved as a distinctly
monophyletic group.
The percentage of trees in which the associated taxa clustered together
is shown next to the branches.
916 ... Naseer, Khalid & Smith
Discussion
Species in the Maculata subclade are characterized in part by the presence of
thin-walled often clavate to pyriform cheilocystidia, phaseoliform spores, and
specific odours, and the stipe base tends to be distinctly bulbous, as found in
I. cookei Bres., I. maculata Boud., and I. quietiodor (Larsson et al. 2009).
Inocybe shawarensis is characterized by its brown fibrillose campanulate
pileus, grayish hymenium, a stipe surface that is silvery white on the lower
two-thirds and dark gray on the upper third, a cylindrical stipe that is slightly
pruinose at apex, slightly bulbous stipe base, an absence of pleurocystidia, and
small phaseoliform yellowish brown spores. Our new species is morphologically
similar to Inocybe quietiodor but separated with 70% bootstrap value from
I. quietiodor (FJ93618, AM882950, AM882961), with which it forms a sister
clade that includes Inocybe cf. reisneri (sampled from Japan).
Compared to I. shawarensis, characterized with a brown campanulate pileus
and gray gills, I. quietiodor has a more prominently umbonate yellowish (5y8/10)
pileus and yellowish lamellae. Inocybe quietiodor is further distinguished
by its yellow cylindrical stipe with the whitish bulbous base and larger and
less phaseoliform spores. Inocybe cookei is distinguished morphologically
by its conspicuously marginate bulbous base (bulb <16 mm) and pyriform
cheilocystidia.
The Indian species I. gregaria K.P.D. Latha & Manim., I. virosa C.K. Pradeep
et al., and I. carnosibulbosa C.K. Pradeep & Matheny falling in the Inosperma
clade “Old World tropical clade 2” may be comparable to our taxon. Inocybe
shawarensis differs from these, however, by its placement in the Maculata
subclade and its smaller basidiomata (Pradeep et al. 2016, Latha & Manimohan
2016).
Acknowledgements
We are sincerely grateful to Dr. Brandon Matheny, Department of Ecology and
Evolutionary Biology, University of Tennessee, Knoxville, USA, for his comments on
an earlier version of this paper. We wish to express our gratitude to Dr. Abdul Rehman
Niazi (Assistant Professor, Department of Botany, University Of the Punjab, Lahore)
for also acting as pre-submission reviewer of this manuscript. We are also thankful to
Dr. Sana Jabeen (University of Education, Lahore) for her help during the manuscript
preparation.
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Ahmad S, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Lahore, Pakistan: Sultan Ahmad
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Anonymous. 1999. District Census Report, Swat. Population census organization, Statistics
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Dentinger BTM, Didukh MY, Moncalvo JM. 2011. Comparing COI and ITS barcode markers for
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Horak E, Matheny PB, Desjardin DE, Soytong K. 2015. The genus Inocybe (Inocybaceae,
Agaricales, Basidiomycota) in ‘Thailand and Malaysia. Phytotaxa 230(3): 201-238.
https://doi.org/10.11646/phytotaxa.230.3.1
Ilyas S, Razaq A, Khalid AN. 2013. Inocybe nitidiuscula and its ectomycorrhizae with Alnus nitida
from Galyat, Pakistan. Mycotaxon 124: 247-254. https://doi.org/10.5248/124.247
Jabeen S, Ahmad I, Rashed A, Khalid AN. 2016. Inocybe kohistanensis, a new species from Swat,
Pakistan. Turkish Journal of Botany 40: 312-318. https://doi.org/10.3906/bot-1501-17
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Kropp BR, Matheny PB, Hutchison LJ. 2013. Inocybe section Rimosae in Utah: phylogenetic
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 919-923
https://doi.org/10.5248/132.919
Phaeomonilia aquatica sp. nov.,
an aquatic hyphomycete from China
JUN-EN HUANG’, HAI-YAN SONG’, XI-GEN HUANG},
JIAN Ma” & D1AN-MinG Hu™
"Jiangxi Key Laboratory for Conservation and Utilization of Fungal Resources,
College of Agronomy, Jiangxi Agricultural University & °College of Science,
Jiangxi Agricultural University, Nanchang, 330045, China
“ CORRESPONDENCE TO: Jian Ma, Dian-Ming Hu, email: hudianming1@163.com
ABSTRACT—A new aquatic hyphomycete, Phaeomonilia aquatica, is described and illustrated
from specimens collected on submerged wood in Jiangxi Province, China. Phaeomonilia
aquatica is morphologically characterized by conidiophores with percurrent proliferation
and aseptate smooth conidia that are spherical, subtriangular, subglobose, ellipsoidal to
subfusiform or obovate and formed in branched acropetal chains. A key to Phaeomonilia
species is provided.
KEY worps — anamorphic fungi, freshwater fungi, lignicolous fungi, taxonomy
Introduction
Hyphomycetous fungi are highly diverse in freshwater habitats (Hu
et al. 2013). As essential components in aquatic ecosystems, they play an
irreplaceable role in the natural cycle of organic materials and energy. Their
global distribution is poorly studied (Shearer et al. 2007), but our group has
investigated the distribution of aquatic hyphomycetes for about ten years
(Hu et al. 2007, 2010a,b, 2012a,b,c, 2013; Huang et al. 2016). The ongoing
investigation in Jiangxi Province, China, has led to the collection of a new
species of Phaeomonilia R.F. Castaneda et al.
Materials & methods
Unidentified wood samples submerged in a freshwater stream were collected in
Jiangxi Province, China, and incubated in moist chambers at room temperature (ca. 25
920 ... Huang & al.
°C). Samples were examined for fungal fruiting bodies using a dissecting microscope.
Observations and photographs were prepared from materials mounted in water and
examined with a Nikon Ni compound microscope (Hu et al. 2012a). The specimens
examined were deposited in the Herbarium of Fungi, Jiangxi Agricultural University,
Nanchang, China (HFJAU).
Taxonomy
Phaeomonilia aquatica J.E. Huang, H.Y. Song, Jian Ma & D.M. Hu, sp. nov. P11
MycoBANkK MB 821142
Differs from Phaeomonilia guangxiensis by its conidiophores with percurrent
proliferations through the apex and by its narrower conidia, and from P. nanningensis
by its conidiophores with percurrent proliferations through the apex and by its shorter
hyaline to yellowish conidia.
TypE—China, Jiangxi Province, Jiujiang, Lushan Mountain, in a stream on submerged
wood, 13 Oct. 2016, J.E. Huang (Holotype, HFJAU 0148).
ETyMOLOGY—aquatica, referring to the aquatic habitat of the fungus.
COLONIES on submerged wood effuse, brown to dark brown, consisting
of scattered single conidiophores over the substrate surface. MYCELIUM
immersed orsuperficial, composed ofbrown, cylindrical, septate, infrequently
branched hyphae. CoNrIDIOPHORES macronematous, mononematous,
sometimes more than 900 um long and 8.5-18 um diam., single, erect,
straight or flexuous, branched or unbranched, smooth, occasionally rough,
15-35-septate, thin-walled, brown to dark brown toward the base, fading
slightly toward the apex, with percurrent proliferations through the apex,
giving rise to new conidiogenous loci at the apex. CONIDIOGENOUS CELLS
holothallic, 12-29 x 8.5-13 um, terminal, subapical or lateral, obovoid,
cupulate or obtrapezoid, smooth, pale brown, with several pale brown to
hyaline trapezoid denticles, 3.0-5.9 x 0.9-2.3 um. Conrp1A thallic-arthric,
1-celled, 8-14 x 7.5-12 um, spherical, subtriangular, subglobose, ellipsoidal
to subfusiform or obovate, hyaline to yellowish, in branched acropetal
chains of about seven conidia, thin-walled, with some small wart on the
surface. Conidial secession schizolytic.
ADDITIONAL SPECIMEN EXAMINED—CHINA, JIANGXI PROVINCE, Jiujiang, Lushan
Mountain, in a stream on submerged wood, 13 Oct. 2016, J.-E. Huang (HFJAU 0323).
PLATE 1. Phaeomonilia aquatica (holotype, HFJAU 0148). a. Conidiophore and conidia
on submerged wood; b-d. Conidiophores and conidia. Note the proliferation (arrowed
in d); e-f. Conidial chains; g. Conidiogenous cell with conidia; h-k. Conidiogenous cells;
l-s. Conidia. Scale bars: a, b = 100 um; c-f = 50 um; g = 20 um; h-s = 5 um.
Phaeomonilia aquatica sp. nov. (China) ... 921
922 ... Huang & al.
Discussion
Castafieda-Ruiz et al. (2007) erected Phaeomonilia for two species,
P. pleiomorpha R.F. Castaneda et al. (type species) and P corticola (R.F.
Castafieda) R.E. Castafieda et al. [= Monilia corticola R.F. Castafeda]. They
compared Phaeomonilia morphologically with similar genera, including
Botryomonilia Goos & Piroz., Staheliella Emden, and Oidiodendron Robak.
Subsequently, Ma et al. (2015) and Yang et al. (2016) described P guangxiensis
Y.R. Ma & X.G. Zhang and P. nanningensis C.L. Yang & X.G. Zhang from China.
Phaeomonilia pleiomorpha was collected from a palm petiole submerged in
a stream, but the other three species were collected from decaying petioles,
twigs, or stems in terrestrial habitats. Phaeomonilia aquatica is the second
species reported from freshwater habitats.
Phaeomonilia aquatica can be easily distinguished from the other
species of Phaeomonilia by its percurrently proliferating conidiophores.
In addition, P pleiomorpha differs from P. aquatica by its smaller, globose,
broadly Y-shaped conidia (4.0-5.8 x 3.9-4.4 um) and its Stylaspergillus-like
synanamorph (Castafeda-Ruiz et al. 2007); P guangxiensis differs by its
slightly shorter and wider, colorless, ellipsoid to globose, broadly Y-shaped
conidia (7-11 x 11-14.5 um; Ma et al. 2015); PR. nanningensis differs by its
longer and slightly wider, brown to pale brown conidia (13.5-25 x 9-13 um;
Yang et al. 2016); and P corticola differs by its narrower, doliiform or ellipsoid
conidia (5.5-16 x 4-5 um; Castaneda-Ruiz et al. 2007). A key to Phaeomonilia
species is presented below.
Key to species of Phaeomonilia
1. Stylaspergillus-like synanamorph present ...................0000 P. pleiomorpha
LANoSivlaspetoriis like synanamorpne, f0 See POR oe oe Le Sree 2
2. Conidia doliiform or ellipsoid, 5.5-16 x 4-5 um... ee ee eee P. corticola
21C conidia ot doliitgtitices A 5-tmTisC ati. the ofc te ee oe ee bey ee by ek eee an ee 3
3. Conidiophores with percurrent proliferation; conidia spherical,
subtriangular, subglobose, ellipsoidal to subfusiform, or obovate. .... P. aquatica
3. Conidiophores without percurrent proliferation; conidia ellipsoidal,
slobose.corn ashaped ce. Meteo oat 5 awe ina ace pig we efag aoe eng ned apn Oo apenas PME age 4
4, Conidia hyaline, 7-11 x 11-14.5 um ....... eee eee eee P. guangxiensis
4, Conidia brown to pale brown, 13.5-25 x 9-13 um ............0006- P. nanningensis
Acknowledgments
The authors express their sincere gratitude to Dr. Eric H.C. McKenzie and Dr.
Huang Zhang for their critical review of the manuscript, to Dr. Shaun Pennycook for
Phaeomonilia aquatica sp. nov. (China) ... 923
nomenclatural review, and to Dr. Lorelei L. Norvell for editorial review. This project
was supported by the National Natural Science Foundation of China (Nos. 31500021,
31460009) and The Key Research and Development Program of Jiangxi Province
(No. 20161 BBF60078).
Literature cited
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Anamorphic fungi from submerged plant material: Phaeomonilia pleiomorpha, P. corticola and
Cacumisporium pleuroconidiophorum. Mycotaxon 100: 327-336.
Hu DM, Zhu H, Cai L, Hyde KD, Zhang KQ. 2007. Sirothecium triseriale, a new chirosporous
anamorphic species from China. Cryptogamie, Mycologie 28: 311-314.
Hu DM, Cai L, Chen H, Bahkali AH, Hyde KD. 2010a. Four new freshwater fungi associated with
submerged wood from Southwest Asia. Sydowia 62: 191-203.
Hu DM, Cai L, Chen H, Bahkali AH, Hyde KD. 2010b. Fungal diversity on submerged wood
in a tropical stream and an artificial lake. Biodiversity and Conservation 19: 3799-3808.
https://doi.org/10.1007/s10531-010-9927-5
Hu DM, Cai L, Bahkali AH, Hyde KD. 2012a. Two new freshwater species of Annulatascaceae from
China. Mycotaxon 120: 81-88. https://doi.org/10.5248/120.81
Hu DM, Cai L, Hyde KD. 2012b. Three new ascomycetes from freshwater in China. Mycologia 104:
1478-1489. https://doi.org/10.3852/11-430
Hu DM, Chen H, Cai L, Bahkali AH, Hyde KD. 2012c. Aquapeziza: a new genus from freshwater,
and its morphological and phylogenetic relationships to Pezizaceae. Mycologia 104: 540-546.
https://doi.org/10.3852/11-123
Hu DM, Liu FE, Cai L. 2013. Biodiversity of aquatic fungi in China. Mycology 4: 125-168.
https://doi.org/10.1080/21501203.2013.835752
Huang JE, Song HY, Ma J, Guan GX, Hu DM. 2016. Xylohyphopsis aquatica sp. nov., a new aquatic
hyphomycete from China. Mycotaxon 131: 391-394. https://doi.org/10.5248/131.391
Ma YR, Xia JW, Zhang XG, Castafieda-Ruiz RF. 2015. New species of Phaeomonilia and Mirandina
from dead branches in China. Mycotaxon 130: 775-781. https://doi.org/10.5248/130.775
Shearer CA, Descals E, Kohlmeyer B, Kohlmeyer J, Marvanova L, Padgett D, Porter D, Raja HA,
Schmit JP, Thorton HA, Voglmayr H. 2007. Fungal biodiversity in aquatic habitats. Biodiversity
and Conservation 16: 49-67. https://doi.org/10.1007/s10531-006-9120-z
Yang CL, Wang JY, Xia JW, Ma YR, Gao JM, Zhang XG. 2016. Phaeomonilia nanningensis sp.
nov. and a new Craspedodidymum record from southern China. Mycotaxon 131: 547-551.
https://doi.org/10.5248/131.547
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 925-931
https://doi.org/10.5248/132.925
Minimelanolocus atlanticus sp. nov.
and M. navicularis from the Brazilian Atlantic Forest
PHELIPE M.O. Costa’, ELAINE MALOsso", MARCELA A. BARBOSA’,
WANDERSON L. TAVARES' & RAFAEL F. CASTANEDA-RUIZ?
"Centro de Biociéncias, Departamento de Micologia &
? Programa de Pés-Graduagao em Biologia de Fungos,
Universidade Federal de Pernambuco, Avenida da Engenharia,
s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil
* Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT)
Alejandro de Humboldt, OSDE, Grupo Agricola,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
* CORRESPONDENCE TO: elaine.malosso@ufpe.br
ABSTRACT—Minimelanolocus navicularis and a new species, M. atlanticus, are described
and illustrated from collections on decaying leaves of unidentified plants. Minimelanolocus
atlanticus is distinguished by oblong, 2-septate conidia, with brown basal and central cells
and gray to subhyaline apical cell and slightly flattened or sub-rounded apex. Notes are
provided on M. navicularis recorded from Brazilian Atlantic Forest.
Key worps—conidial fungi, asexual fungi, neotropic, systematic
Introduction
The genus Minimelanolocus, established by Castafieda-Ruiz et al. (2001)
and typified by Pseudospiropes navicularis R.F. Castaneda [= M. navicularis],
is characterized by conidiophores that are macronematous, mononematous,
erect, unbranched, smooth or verrucose and conidiogenous cells that
are polyblastic, sympodial elongated, terminal, indeterminate, and with
inconspicuous or slightly prominent, narrow, opaque, refractive to somewhat
obscure conidiogenous loci. Minimelanolocus currently comprises 31 species
(Castaneda-Ruiz et al. 2001, 2003; Fiuza et al. 2017; Heredia et al. 2014;
Hernandez-Restrepo et al. 2012; Liu et al. 2015; Ma et al. 2008, 2011; Tian
926 ... Costa & al.
et al. 2016; Xia et al. 2014; Zhang et al. 2009, 2010), most of which occur as
saprobes on decaying leaves, rotten wood, dead branches, bamboo culms, and
submerged plant debris in aquatic habitats.
During a mycological survey of microfungi associated with leaf litter in a
Brazilian Atlantic forest, two conspicuous fungi representing Minimelanolocus
were collected. We describe one as a new species and present a new description
of the other, M. navicularis.
Materials & methods
Individual collections were placed in plastic bags, taken to the laboratory, and
treated according to Castafeda-Ruiz et al. (2016). Mounts were prepared in lactic
acid (90%) or in polyvinyl alcohol-glycerol (8 g PVA in 100 ml H,O + 5 ml glycerol)
and lactofuchsin (0.1 g acid fuchsin in 100 ml 85% lactic acid) following Carmichael
(1955). Measurements were made at a magnification of x1000 under a Nikon Eclipse
Ni-U microscope with bright field optics, and photomicrographs were taken using DIC
optics with a Nikon DS-Fi2 camera. The holotype was deposited in the Herbarium of
Universidade Federal de Pernambuco, Recife, Brazil (URM).
Taxonomy
Minimelanolocus atlanticus PM.O. Costa, Malosso &
R.F. Castafieda, sp. nov. Figs 1, 2
INDEX FUNGORUM IF 554056
Differs from Minimelanolocus obscurus by its 2-septate and versicolored conidia.
Type: Brazil, Pernambuco, Igarassu, RPPN Reftigio Ecoldgico Charles Darwin, 7°48’S
34°57’W, on decaying leaves of an unidentified plant, 12.V.2017, coll. RM.O. Costa
(Holotype, URM 90133).
EryMo_oey: Latin, atlanticus, refers to habitat of the fungus, Brazilian Atlantic Forest.
CoLonies on the natural substrate, effuse, hairy, brown. Mycelium superficial
and immersed, composed of septate, branched, brown, smooth-walled
hyphae, 2-3 um diam. CONIDIOPHORES macronematous, mononematous,
erect, straight or flexuous, cylindrical, slightly geniculate near the apex,
unbranched, 7-15-septate, dark brown below, pale brown toward the apex,
200-300 x 5-7 um, smooth. CONIDIOGENOUS CELLS polyblastic, somewhat
sinuate with inconspicuous denticulate or faintly cicatrized, opaque loci,
terminal, integrated, indeterminate with several sympodial extensions, pale
brown, 30-60 x 3.5-4 um. Conidial secession schizolytic. Conrp1a solitary,
acropleurogenous, oblong, sub-rounded or slightly flattened at the apex,
2-septate, bicolor with basal and central cells brown to dirty brown, apical
cell smoked-gray to subhyaline, smooth, dry, 18-21 x 4-5 um. Sexual state
unknown.
Minimelanolocus atlanticus sp. nov. (Brazil) ... 927
Fic. 1. Minimelanolocus atlanticus (ex holotype URM 90133). A. Conidia; B. Conidiophores
and conidiogenous cells; C. Conidiogenous cells; D. Conidiogenous cells and conidia.
C & D share the same scale.
928 ... Costa & al.
Fic. 2. Minimelanolocus atlanticus (ex holotype URM 90133).
Conidiogenous cells and conidia.
Minimelanolocus atlanticus sp. nov. (Brazil) ... 929
Notes: Minimelanolocus obscurus (Matsush.) R.E Castafieda & Heredia [=
Pseudospiropes obscurus Matsush.] is similar to M. atlanticus in producing
oblong conidia but is distinguished by conidia that are 1-4-septate (usually
3-septate), larger (18-34 x 4.5-8 um), and uniformly colored subhyaline or
pale brown (Castafeda-Ruiz et al. 2001, Liu et al. 2015, Matsushima 1983).
Minimelanolocus navicularis (R.F. Castafieda) R.E Castafieda,
Cryptog. Mycol. 22: 9 (2001). Fic. 3
CONIDIOPHORES macronematous, mononematous, erect, straight or
flexuous, unbranched, cylindrical, inflated at the base, 2-5-septate, smooth,
brown, pale brown above, smooth, 43-70 x 4-6 um. CONIDIOGENOUS
CELLS polyblastic, integrated, terminal, indeterminate, sympodial elongated,
pbebe bee
10 pm
10 pm
Fic. 3. Minimelanolocus navicularis (ex URM 90185).
A. Conidia; B. Conidiophores and conidiogenous cells.
930 ... Costa & al.
inconspicuous denticulate, slightly melanized at the loci, 14-22 x 3-4 um, pale
brown. Conidial secession schizolytic. Conrp1a solitary, acropleurogenous,
navicular, straight, smooth, 3-euseptate, 17-24 x 5-8.5 um, obscure at the septa,
dry, with subhyaline to pale brown end cells, brown to dark brown central cells.
SPECIMEN EXAMINED: BRAZIL, PERNAMBUCO STATE, Lagoa da Mata, Ilha de
Itamaraca, 7°46’S 34°51’W, on decaying leaves of an unidentified plant, 23.X1.2016, coll.
W.L. Tavares (URM 90185).
Notes: Minimelanolocus navicularis has been previously recorded in Brazil
by Santa Izabel et al. (2011); their specimen had longer conidiophores
(45-155 um) than our specimen, but the conidia in both specimens closely
match those of the type species (20-25 x 6-8 um; Castaneda-Ruiz 1987).
Acknowledgments
The authors express their sincere gratitude to Dr. Xiu-Guo Zhang and Dr. De-Wei Li
for their critical review of the manuscript. The authors are grateful to the Coordenacao
de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) (Project 88881.062172/
2014-01) and Programa Ciéncia sem Fronteiras for financial support. RFCR is grateful
to the Cuban Ministry of Agriculture and Programa de Salud Animal y Vegetal (Project
P131LH003033) for facilities. We acknowledge the facilities provided by Dr. P.M.
Kirk and Drs. V. Robert and K. Bensch, through the Index Fungorum and MycoBank
websites. Dr. Lorelei Norvell’s editorial review and Dr. Shaun Pennycook’s nomenclature
review are greatly appreciated.
Literature cited
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Castaneda-Ruiz RE. 1987. Fungi cubenses II. La Habana, Cuba. 22 p.
Castaneda-Ruiz RE Heredia G, Reyes M, Arias RM, Decock C. 2001. A revision of
the genus Pseudospiropes and some new taxa. Cryptogamie, Mycologie 22: 3-18.
https://doi.org/10.1016/S0181-1584(01)01057-0
Castafieda-Ruiz RF, Guarro J, Velasquez-Noa S, Gené J. 2003. A new species of Minimelanolocus
and some hyphomycete records from rain forests in Brazil. Mycotaxon 85: 231-239.
Castafieda-Ruiz RF, Heredia G, Gusmao LFP, Li DW. 2016. Fungal diversity of Central and
South America. 197-217, in: DW Li (ed.). Biology of Microfungi. Springer International
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Fiuza PO, Conceigéo LB, Marques MFO, Gusmao LFP, Rafael F. Castafieda-Ruiz RF. 2017.
Dictyotrichocladium aquaticum gen. & sp. nov. and Minimelanolocus aquatilis sp. nov.
from freshwater in Brazil's semiarid region. Mycotaxon 132(2): 433-440. https://doi.
org/10.5248/132.433.
Heredia G, Arias RM, Castafieda-Ruiz RF, Minter DW. 2014. New species of Lobatopedis and
Minimelanolocus (anamorphic fungi) from a Mexican cloud forest. Nova Hedwigia 98: 31-40.
https://doi.org/10.1127/0029-5035/2013/0146
Hernandez-Restrepo M, Castafieda-Ruiz RF, Gené J, Guarro J, Minter DW, Stadler M. 2012.
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pediculata comb. nov. Mycotaxon 122: 135-143. https://doi.org/10.5248/122.135
Minimelanolocus atlanticus sp. nov. (Brazil) ... 931
Liu XY, Udayanga D, Luo ZL, Chen LJ, Zhou DQ, Hyde KD. 2015. Backbone tree for Chaetothyriales
with four new species of Minimelanolocus from aquatic habitats. Fungal Biology 119:
1046-1062. https://doi.org/10.1016/j.funbio.2015.08.005
Ma J, Zhang YD, Ma LG, Zhang XG. 2008. Two new species of the Minimelanolocus in China.
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Ma J, Zhang YD, Ma LG, Zhang XG. 2011. Two new Minimelanolocus species from southern China.
Mycotaxon 117: 131-135. https://doi.org/10.5248/117.131
Matsushima T. 1983. Matsushima mycological memoirs, no. 3. 90 p.
Santa Izabel TS, Santos DS, Almeida DAC de, Gusmao LFP. 2011.Fungos conidiais do bioma
Caatinga II. Novos registros para o continente americano, Neotrépico, América do Sul e
Brasil. Rodriguésia 62(2): 229-240. https://doi.org/10.1590/2175-7860201162201
Tian Q, Doilom M, Luo ZL, Chomnunti P, Bhat JD, Xu JC, Hyde KD. 2016. Introducing
Melanoctona tectonae gen. et sp. nov. and Minimelanolocus yunnanensis sp. nov.
(Herpotrichiellaceae, Chaetothyriales). Cryptogamie, Mycologie 37: 477-492.
https://doi.org/10.7872/crym/v37.iss4.2016.477
Xia JW, Ma LG, Castafieda-Ruiz RF, Zhang XG. 2014. Minimelanolocus bicolorata sp. nov.,
Paradendryphiopsis elegans sp. nov. and Corynesporella bannaense sp. nov. from southern
China. Mycoscience 55: 299-307. https://doi.org/10.1016/j.myc.2013.11.003
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Mycotaxon 114: 373-376. https://doi.org/10.5248/114.373
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 933-941
https://doi.org/10.5248/132.933
New records of Amanita from
Tehuacan-Cuicatlan Biosphere Reserve, México
EVANGELINA PEREZ-SILVA* & ABRAHAM J. MEDINA-ORTIZ
Botany Department, Laboratory of Mycology,
Institute of Biology, National Autonomous University of Mexico (UNAM),
University City, C.P. 04510, México City, México
* Correspondence to: psilva@ib.unam.mx
ABSTRACT—Specimens of Amanita caesarea, A. flavoconia, A. flavorubescens, A. fulva,
A. onusta, A. pantherina, A. rubescens, A. vaginata, and A. verna are described as new records
from Tehuacan-Cuicatlan Biosphere Reserve. There were no records of mycetism produced
by the consumption of those species in the reserve.
Key worps—Amanitaceae, Agaricales, edible, Oaxaca, taxonomy, toxic
Introduction
Ruiz-Oronoz & Herrera (1948) and Herrera (1950) were to first to initiate
mycological research in the Cuicatlan district of Oaxaca. In 1998, President
Ernesto Zedillo Ponce de Leon proclaimed the studied region as the Natural
Protected Area of Tehuacan-Cuicatlan Biosphere Reserve in the states of Puebla
and Oaxaca, México, with the intent of conserving its plant, animal, and fungal
diversity as well as its huge human cultural wealth. Since mycological studies
were restarted in 1996, fungi have been collected in different the vegetation
types of tropical subdeciduous forest, deciduous forest, and pine-oak forest.
As a result of collaborative work amongst the Institute of Biology (National
Autonomous University of Mexico), Ejidal and Communal Commission of
Tehuacan-Cuicatlan, and National Commission of Natural Protected Areas
with the communities of San Pedro Jocotipac and Santa Maria Papalo, nine
species of Amanita (Agaricales, Amanitaceae) have been recorded from this
region for the first time.
934 ... Pérez-Silva & Medina-Ortiz
The aim of this research was to extend the taxonomic knowledge of Amanita
as an exploitable natural resource. Of the nine Amanita spp. identified, four
were edible and five were toxic (one fatally poisonous). In view of the fact
that the reserve’s mycobiota is basically unknown, it is imperative that fungal
research be continued in the region.
Materials & methods
Specimens were collected in oak forests in the Cuicatlan district. Digital pictures,
field data, and macro- and microscopical characterization of samples were performed.
Collections are deposited in the Fungus Collection of the National Herbarium, Institute
of Biology, National Autonomous University of México (MEXU). Colour codes in
descriptions follow Kornerup & Wanscher (1978). Morphological structures were
observed using a stereomicroscope and compound light microscope. Basidiomes were
sectioned and mounted in ammonia water (1%), KOH (5%), Melzer reagent, neutral
red (1%), and cotton blue in lactic acid solutions. Species were identified consulting Bas
(1969), Jenkins (1977), Arora (1986), Pérez-Silva & Herrera (1991), Phillips (1991), and
Tulloss et al. (1992).
Taxonomy
Amanita caesarea (Scop.) Pers., Syn. Meth. Fung. Pars 2: 252 (1801). Fie. 1
BASIDIOME 90-120 mm high. Prteus 5.5-10.5 mm diam., convex, becoming
flatter in age, with long striate margin; disc high red to vivid red (11A8), paler
toward the margin. LAMELLAE 8 mm broad, adnate, with entire edge, pale
yellow (4A3). Stipe 9-12 x 8-17 cm, pale yellow, with darker orange fiber or
patches; annulus pendulous, membranous, yellowish, attached to the upper
stem, often striate; no basal bulb, white volval sac. CONTEXT white to yellowish.
ODOR & TASTE pleasant.
BASIDIOSPORES 8-16.5 x 5.5-8 um, elliptical, thin-walled; basal cylindric
apiculus; uniguttulate, broadly ellipsoid; hyaline, inamyloid. BAsip1A 38.5-49.5
x 10-11 um, tetrasporic, with basal clamp connections.
EcoLoGy—Solitary (rarely in groups) in pine-oak woods during July-
November; mycorrhizal; widely distributed in México, where it is reported
as an edible of excellent culinary value; commonly called “yellow fungus”
(Pérez-Silva & Herrera 1991).
SPECIMENS EXAMINED: MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F Medina Ruiz, A. de la Cruz
Martinez (MEXU 27820); El Sdétano, 11 Jul 2014, E. Pérez-Silva, A.J. Medina-Ortiz,
A. Garcia-Mendoza, F. Hernandez (MEXU 27827).
COMMENTS—Amanita caesarea is easily distinguished by its bright yellow gills
and stipe, brilliant red to orange cap with striate margin, and voluminous white
Amanita spp. new from Tehuacaén-Cuicatlan ... 935
FiGurREs 1-8. Amanita species basidiomata. 1: A. caesarea (MEXU 27827). 2: A. flavoconia (MEXU
27828). 3: A. fulva (MEXU 27833). 4:.A. onusta (MEXU 27823). 5:.A. pantherina (MEXU 27835).
6: A. rubescens (MEXU 27832). 7: A. vaginata (MEXU 27824). 8: A. verna (MEXU 27825).
volva (Arora 1986). Fresh mushrooms are sold at markets and along roadsides
in México (Montoya et al. 2001) and the United States (Volk 2017). Amanita
jacksonii is similar, but differs by its larger, bright orange-red basidiome and
occurrence in northern mixed forests of Canada.
Amanita flavoconia G.F. Atk., J. Mycol. 8(3): 110 (1902). Fic. 2
BASIDIOME 80 mm high. Piteus 40-60 mm diam., convex, becoming
broadly flatter when maturing, yellowish red (8B8); surface often viscid, with
bright yellow universal veil remnants; not striate margin. LAMELLAE 7 mm
broad, white to pale yellow (4A3), free from stem, with entire edge. STIPE 60-80
x 5-9 mm, white to yellowish, membranous annulus to the apex, pale yellow;
swollen basal bulb, 20 mm diam., covered with yellow floccose veil fragments,
friable volva. CONTEXT white. ODOR & TASTE pleasant.
BASIDIOSPORES 8-9 xX 6-7 um, ovoid, hyaline, amyloid, thin-walled.
BASIDIA 33-38 x 8-8.5 um, clavate, tetrasporic. PILEIPELLIS hyphae 2-5 um
wide, gelatinized filamentous; oleiferous hyphae 2-4 um wide.
936 ... Pérez-Silva & Medina-Ortiz
EcoLtocy—Solitary in pine-oak woods; fructification during July-
November; wide distribution in México (Pérez-Silva & Herrera 1991).
Villanueva-Jiménez et al. (2006) cited from Ixtlan, Oaxaca.
SPECIMEN EXAMINED: MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F. Medina-Ruiz, A. de la Cruz
Martinez (MEXU 27821); El Sdétano, 11 Jul 2014, E. Pérez-Silva, A.J. Medina-Ortiz,
A. Garcia-Mendoza, FE Hernandez (MEXU 27828).
CoMMENTS—Amanita flavoconia has been reported as toxic in several localities
in USA (Metzler & Metzler 1992) and may cause gastrointestinal mycetism.
Benedict et al. (1966) did not detect ibotenic acid or muscimol nor did Chilton
& Ott (1976) find toxic metabolites. Aroche et al. (1984) reported presence of
amanitins.
Amanita flavorubescens G.F. Atk., J. Mycol. 8(3): 111 (1902).
BASIDIOME 60 mm high. PrLrEus 40-60 mm diam., convex, becoming flatter
when maturing, orange yellow (4A8); thick yellow warts of volva around center.
LAMELLAE 5 mm broad, white to pale yellow (4A3), free from stipe, white
pulverulent edge. Stipe 50-70 x 3-7 mm, yellow to pale yellow, narrower at
the apex, pruinose to powdery; annulus membranous, skirt-like; basal bulb,
5-15 mm diam., with yellowish margin. CONTEXT white, reddish at the base.
ODOR & TASTE not distinctive.
BASIDIOSPORES 9-10 x 5-7 um, ellipsoid, thin-walled, uniguttulate, with
basal eccentric apiculus, amyloid. Basrp1a tetrasporic. PILEIPELLIS hyphae
2-5 um wide, filamentous; oleiferous hyphae present.
EcoLtoGy—Solitary in pine-oak woods during July-November; mycorrhizal;
widely distributed in Chihuahua, Estado de México, Hidalgo and Morelos
States of México (Pérez-Silva & Herrera 1991). Edibility unknown.
SPECIMEN EXAMINED: MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F. Medina Ruiz, A. de la Cruz
Martinez (MEXU 27822); Santa Maria Papalo, 12 Jul 2014, E. Pérez-Silva, A.J. Medina-
Ortiz, A. Garcia-Mendoza, EF. Hernandez (MEXU 27829).
ComMMENTsS—Amanita flavorubescens is easily recognized by its striking
yellow pileus with yellow warts, free white crowded lamellae with floccose
edges, yellowish stipe with a superior annulus and slightly reddish persistently
bulbous base.
Amanita fulva Fr., Observ. Mycol. 1: 2 (1815). Fic. 3
BASIDIOME 90 mm high. PiLEus 25-45 mm diam., oval, becoming broadly
convex or flat in age; brownish grey (8D2) to dark brown (8F5); surface viscid
Amanita spp. new from Tehuacan-Cuicatlan ... 937
when moist; remarkably striate margin; LAMELLAE 4 mm broad, whitish (9A1),
free from the stem. STIPE 95 x 5 mm, with very fine hairs, without annulus; the
base enclosed by a white large volva. CONTEXT white. ODOR & TASTE indistinct.
BASIDIOSPORES 8-10 x 9 um, globose, apiculus basal and eccentric, hyaline,
inamyloid. Basip1a 23-38 x 5-7 um, tetrasporic. PILEIPELLIS hyphae 2-8 um
wide, thin filamentous, brown.
EcoLocy—Solitary in pine-oak woods during July; in México widely
distributed in Durango, Estado de México, Hidalgo, Morelos, Puebla and
Veracruz states (Pérez-Silva & Herrera 1991). Edible, although toxic when raw
or poorly cooked (Pérez-Silva 2004).
SPECIMEN EXAMINED: MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
El Sétano, 11 Jul 2014, E. Pérez-Silva, A.J. Medina-Ortiz, A. Garcia-Mendoza,
FE Hernandez (MEXU 27830); Santa Maria Papalo, 11 Jul 2014, E. Perez-Silva,
A.J. Medina-Ortiz, A. Garcia-Mendoza, F. Hernandez (MEXU 27833).
ComMMENTS—Amanita fulva is easily recognized by its often umbonate pileus,
striate margin, free whitish lamellae, the stipe lacking an annulus, large white
volva, and inamyloid spores.
Amanita onusta (Howe) Sacc., Syll. Fung. 9: 1 (1891). Fic. 4
BASIDIOME 90-180 mm high. PiLeus 45-90 mm diam., convex, expanding
to flat in age; white (10E2) to pale gray (1B1), covered with dark grey to
brownish grey pyramidal warts, 5 mm high (but shorter toward margin),
veil fragments frequent. LAMELLAE 10 mm broad, white to pale gray (1B1),
ventricose, adnexed, floccose. STIPE 90-180 x 15-25 mm, brownish-grey,
paler towards the top; annulus apical but lost in age; base a ventricose to
napiform basal bulb, 25-35 mm diam.; volva floccose, fibrillose, brownish
grey, remnants in circles around stipe. CONTEXT whitish. ODOR & TASTE
unpleasant.
BASIDIOSPORES 9-10 x 6-7 um, ellipsoid, thin-walled, amyloid. BAsrp1a
38-44 x 6-9 um, pyriform; clamp connections present. PILEIPELLIS thick
layer interwoven, hyphae 3 um diam.; oleiferous hyphae abundant.
EcoLocy—Solitary in pine-oak woods during November; mycorrhizal;
previously reported in México from Estado de México and Veracruz State
(Pérez-Silva & Herrera, 1991) and now for the first time in Oaxaca State.
Little is known regarding its edibility, but its unpleasant odor probably deters
consumption.
SPECIMEN EXAMINED: MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F. Medina Ruiz, A. de la Cruz
Martinez (MEXU 27823).
938 ... Pérez-Silva & Medina-Ortiz
COMMENTS—Amanita onusta is characterized by its white to pale grey pileus
covered with dark gray to brownish grey pyramidal warts, its dirty white stipe
with a paler apex, and elliptical amyloid spores.
Amanita pantherina (DC.) Krombh.,
Naturgetr. Abbild. Schwamme 4: t.29 f. 10-13 (1836). Fic. 5
BASIDIOME 50-150 mm high. Piteus 50-70 mm diam., convex, finally
flattened; brownish grey (9D3), surface slightly viscid, covered with adnate
white warts. LAMELLAE 8 mm broad, white, free, narrowed to the stem. STIPE
80-120 x 20-50 mm, white, apical annulus, smooth; basal bulb, volva with 2-3
concentric rings. CONTEXT white. ODOR & TASTE unpleasant.
BASIDIOSPORES 7-10 x 6-9 um, ovoid, thin-walled, with hilar appendix.
BASIDIA 38-55 x 10-15 um.
EcoLocy—Solitary in pine-oak woods during July; reported from
Chihuahua, Hidalgo, Estado de México, and Puebla states (Pérez-Silva &
Herrera 1991) and from Ixtlan Mountain range in Oaxaca State by Villanueva-
Jiménez et al. (2006). Although there is no data regarding intoxication from
the Cuicatlan District, Amanita pantherina is known to produce ibotenic
acid (Pérez-Silva 2004; Pérez-Silva et al. 2006, 2008), which causes nervous
mycetism.
STUDIED MATERIAL. MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
El Sotano, 11 Jul 2014, E. Pérez-Silva, A.J. Medina-Ortiz, A. Garcia-Mendoza,
FE Hernandez (MEXU 27831); Santa Maria Papalo, 12 Jul 2014, E. Pérez-Silva,
A.J. Medina-Ortiz, A. Garcia-Mendoza, F. Hernandez (MEXU 27835).
CoMMENTS—Amanita pantherina is easily recognized by its viscid brownish
pileus decorated with white to cream patches and initially striate curved
margin, free white lamellae, white stipe with a superior annulus and basal bulb,
and its 2-3 concentrically ringed volva.
Amanita rubescens Pers., Tent. Disp. Meth. Fung.: 67 (1797). FIG. 6
BASIDIOME 60-80 mm high. Pireus 40-85 mm diam., convex, becoming
flat; reddish to reddish white (13A2); surface covered with adnexed warts;
spotted reddish where injured. LAMELLAE 4 mm broad, white to pale reddish
(13A2), free. StrpE 70-100 x 10-25 mm, concolorous with the pileus, white;
annulus large, membranous; basal bulb, staining reddish; volva scaly, reddish
grey (7B2), sometimes leaving remnants in circles. CONTEXT white, staining
pinkish when injured. ODOR & TASTE pleasant.
BASIDIOSPORES 8-9 x 5-7 um, elliptical, thin-walled, smooth, amyloid.
Basip1a 44-47 x 8 um, cylindrical, tetrasporic.
Amanita spp. new from Tehuacan-Cuicatlan ... 939
EcoLocy—Solitary in pine-oak woods during July, widely distributed
in México from Baja California, Chihuahua, Distrito Federal, Durango,
Hidalgo, Jalisco, Michoacan, Morelos, Nuevo Leén, Oaxaca, Puebla, Sonora,
Veracruz, and Zacatecas states (Pérez-Silva & Herrera 1991). Considered
edible after discarding the first cooking water, which removes the toxic protein
rubescenslysine. Nonetheless, Amanita rubescens is not usually consumed
when parasitized by Hypomyces hyalinus.
STUDIED MATERIAL. MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
El Sétano, 11 Jul 2014, E. Pérez-Silva, A.J. Medina-Ortiz, A. Garcia-Mendoza,
F. Hernandez (MEXU 27832).
CoMMENTS—Amanita rubescens is easily recognized by its white to reddish
pileus.
Amanita vaginata (Bull.) Lam., Encycl. Méth. Bot.. 1(1): 109 (1783). Fig. 7
BASIDIOME 70 mm high. PrLEus 30 mm diam., ovoid, becoming flat with a
low umbo; brownish grey (7E2), viscid when wet, covered with membranous
white universal veil patches; margin radially striate. LAMELLAE 5 mm broad,
close, free, white to pallid (1A2). Stipe 60-70 mm long, white, with scales;
base enclosed in a sac-like volva, membranous, white. CONTEXT white. ODOR
& TASTE pleasant.
BASIDIOSPORES 10-11 um, globose, hyaline, inamyloid. Basrp1a 50-60 um,
tetrasporic. PILEIPELLIS with filamentous cuticle and abundant oleiferous
hyphae.
EcoLocy—Solitary in pine-oak woods during November; widely distributed
in México from Chihuahua, Estado de México, Hidalgo, and Oaxaca states
(Pérez-Silva & Herrera 1991). Considered edible.
STUDIED MATERIAL. MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F Medina-Ruiz, A. de la Cruz-
Martinez (MEXU 27824).
COMMENTS—Amanita vaginata contains hemolysins and may cause severe
gastrointestinal mycetism when consumed raw. Pérez-Silva & Herrera (1991)
cite several common names for this species.
Amanita verna (Bull.) Lam., Encycl. Méth. Bot. 1(1): 113 (1783). Fic. 8
BASIDIOME 100 mm high. Prreus 70-90 mm diam., hemispheric with
flattened center, becoming applanate; white (23A1) to whitish cream
(24A1), occasionally with some scales. LAMELLAE 4-5 mm broad, white
(24A1), free, edges serrate, floccose. STIPE 80-90 x 40-50 mm, cylindrical,
white, pale yellow in 10% KOH; annulus superior, membranous, fragile;
9AO ... Pérez-Silva & Medina-Ortiz
basal bulb 40-50 x 30-50 mm, covered with a persistent membranous
saccate volva. CONTEXT white. ODOR & TASTE unpleasant. EDIBILITY
poisonous.
BASIDIOSPORES 9-10 x 6-8 um, ellipsoid, thin-walled, hyaline, amyloid.
Basip1A 40-44 x 5-6 um, tetrasporic, yellowish.
EcoLtocy—Solitary in pine-oak woods during November; widely
distributed in México from Durango, Estado de México, Guanajuato,
Guerrero, Hidalgo, Michoacan, Morelos, Sonora, and Veracruz states
(Pérez-Silva & Herrera 1991); previously reported from Ixtlan belt of
Oaxaca State (Villanueva-Jiménez et al. 2006).
STUDIED MATERIAL. MEXICO. Oaxaca: Cuicatlan District, San Pedro Jocotipac,
Ciénega de Cochino, 17 Nov 2013, A.J. Medina-Ortiz, F. Medina Ruiz, A. de la Cruz
Martinez (MEXU 27825); Santa Maria Papalo, 12 Jul 2014, E. Pérez-Silva, A.J. Medina-
Ortiz, A. Garcia-Mendoza, EF Hernandez (MEXU 27834).
COMMENTS—Amanita verna is similar to A. virosa Bertill., which differs
in its positive reaction in 5% KOH and ovoid amyloid spores. Extremely
toxic, A. verna frequently causes fatalities and cyclopeptide mycetism due to
amanitins and other toxins; commonly known as “crazy fungus” (Pérez-Silva
2004; Pérez-Silva et al. 2006, 2008). Bresinsky & Besl (1990) cite the species
as mycorrhizal.
Discussion
Five of the nine Amanita species reported here are toxic, causing
cyclopeptide mycetism (A. verna), pantherina mycetism (A. pantherina),
and possibly a type of gastrointestinal mycetism when eaten raw
(A. flavoconia, A. flavorubescens, and A. vaginata). Amanita caesarea
is the only wild species with excellent cooked or raw edibility, although
A. rubescens can be safely eaten after its boiling water is discarded. The
edibility/toxicity of A. onusta remains unknown.
All nine amanitas are reported for the first time from Tehuacan-Cuicatlan
biosphere reserve, which extends their known Mexican distribution.
Acknowledgments
This article is dedicated to Dr. Teofilo Herrera, Emeritus Researcher of UNAM, for
his outstanding academic trajectory and forming mycologists in Mexico. We thank SNI
for financial support, Dr. Martin Esqueda and Dr. Gabriel Moreno for pre-submission
expert reviews, Dr. Shaun R. Pennycook for critical reading of the manuscript and
suggestions, MSc. C.E. Aguirre-Acosta for curatorial support in MEXU, and David
Esqueda for translation.
Amanita spp. new from Tehuacan-Cuicatlan ... 941
Literature cited
Aroche M, Cifuentes J, Lorea FG, Bonavides J, Garcia H. 1984. Macromicetos toxicos y
comestibles de una regi6n comunal del Valle de México. Boletin de la Sociedad Mexicana
de Micologia 19: 291-318.
Arora D. 1986. Mushrooms demystified: a comprehensive guide to the fleshy fungi. Ten Speed
Press. Berkeley. USA.
Bas C. 1969. Morphology and subdivision of Amanita and a monograph of its section Lepidella.
Persoonia 5: 285-579.
Benedict R, Tyler Jr V, Brady L. 1966. Chemotaxonomic significance of isoxazole derivatives in
Amanita species. Lloydia 29: 333-341.
Bresinsky A, Besl H. 1990. A colour atlas of poisonous fungi: a handbook for pharmacists, doctors,
and biologist. Woolfe, London.
Chilton WS, Ott J. 1976. Toxic metabolites of Amanita pantherina, A. cothurnata, A. muscaria and
other Amanita species. Lloydia 39: 150-157.
Herrera T. 1950. Un hongo interesante en la regidn de Cuicatlan, Oaxaca. Anales del Instituto de
Biologia, Universidad Nacional Aut6noma de México 21: 17-21.
Jenkins DT. 1977. A taxonomic and nomenclatural study of the genus Amanita Section Amanita for
North America. Bibliotheca Mycologica Band 57. J. Cramer, Stuttgart, Germany.
Kornerup A, Wanscher JH. 1978. Methuen handbook of color. Methuen Co. London. UK.
Metzler S, Metzler V. 1992. Texas mushrooms a field guide. University of Texas. Austin. USA.
Montoya A, Estrada A, Kong A, Juarez-Sanchez L. 2001. Commercialization of wild mushrooms
during market days of Tlaxcala, Mexico. Micologia Aplicada Internacional 13(1): 31-40.
Pérez-Silva E. 2004. Intoxicacién por hongos. 305-309, in: OB Martinez Pantaleén (ed.).
Intoxicaciones. Temas de Pediatria. Asociacién Mexicana de Pediatria AC. McGraw-Hill.
Interamericana Editores, S.A. de C.V. México.
Pérez-Silva E, Herrera T. 1991. Iconografia de macromicetos de México. I Amanita. Instituto de
Biologia, Universidad Nacional Autonoma de México.
Pérez-Silva E, Esqueda M, Herrera T. 2008. Macromicetos téxicos de Sonora, México. Revista
Mexicana de Micologia 28: 81-88.
Pérez-Silva E, Esqueda M, Herrera T, Coronado M. 2006. Nuevos registros de Agaricales de Sonora,
México. Revista Mexicana de Biodiversidad 77: 23-33.
Phillips R. 1991. Mushrooms of North America. Little Brown and Co. New York. USA.
Ruiz-Oronoz M, Herrera T. 1948. Levaduras, hongos macroscdépicos, liquenes y hepaticas
colectados en Cuicatlan. Oaxaca. Anales del Instituto de Biologia, Universidad Nacional
Auténoma de México 19: 299-316.
Tulloss RE, Ovrebo CL, Halling RE. 1992. Studies on Amanita (Amanitaceae) from Andean
Colombia. Memoirs of the New York Botanical Garden 66. 46 p.
Villanueva-Jiménez E, Villegas-Rios M, Cifuentes-Blanco J, Leén-Avendafio H. 2006.
Diversidad del género Amanita en dos areas con diferente condicion silvicola en Ixtlan de
Juarez, Oaxaca, México. Revista Mexicana de Biodiversidad 77: 17-22.
Volk TJ. 2017. Amanita caesarea. TomVolkFungi.net. Accessed January/2017:
http://botit.botany.wisc.edu/toms_fungi/mar2002.html
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 943-950
https://doi.org/10.5248/132.943
Thelephora iqbalii sp. nov. from the
Himalayan moist temperate forests
of Pakistan
ABDUL NASIR KHALID’ & MUHAMMAD HANIkE”?
' Department of Botany, University of the Punjab,
Quaid-e-Azam Campus, Lahore, 54590, Pakistan
? Department of Botany, GC University, Lahore, Pakistan
* CORRESPONDENCE TO: dr.mhanif@gcu.edu.pk
ABSTRACT—A new thelephoroid species, Thelephora iqbalii, collected from the Himalayan
moist temperate (HMT) forests of Pakistan, is characterized by small coralloid basidiomata
bearing long, narrow ridges with fine hairs on their branches and hymenia that are cyanescent
in KOH. An ITS-based phylogeny comparing T. iqbalii with other Thelephora species is also
presented.
Key worps—Basidiomycota, DNA, Thelephoraceae, angular lobate basidiospores
Introduction
The genus Thelephora Ehrh. ex Willd. is cosmopolitan (Corner 1968;
Stalpers 1993; Kdljalg 1996; Maas Geesteranus 1971, 1975) and represented by
52 species worldwide (Kirk et al. 2008, Ramirez-Lopez et al. 2017, Vizzini et al.
2016). Its basidiomata are distinguished by their pileate and stipitate/sessile or
sometimes resupinate forms (Ahmad 1972).
Thelephora basidiomata range in shape from coral-like to distinctly pileate
(Corner 1968). However, morphology alone is not always sufficient for
correct identification. Recent molecular analyses have contributed to a better
understanding of basidiomycete diversity (Gardes & Bruns 1993) and ITS-
nrDNA primers are widely being used for this purpose (EI Karkouri et al. 2005,
2006; Gardes & Bruns 1993; Horton 2002).
944 ... Khalid & Hanif
The Himalayan moist temperate forests (HMT) have been previously
included among 25 biodiversity hotspots in Indo-Burma (Myers et al.
2000), from which they were separated and designated as independent
among the 34 currently recognized hotspots of biodiversity (Mittermeier
et al. 2011).
Eight Thelephora species have previously been reported from Pakistan:
T. anthocephala (Bull.) Fr., T: arbuscula Corner, T: caryophyllea (Schaeff.)
Pers., T: fucoides Corner, T: palmata (Scop.) Fr., T. penicillata (Pers.) Fr.,
and T: terrestris Ehrh. (Ahmad 1956, 1972; Ahmad et al. 1997). Below we
describe a new thelephoroid species from the HMT forests of Pakistan
based on morphological and molecular evidence.
Materials & methods
TABLE 1. Thelephora rDNA sequences used in phylogenetic analyses
TAXON ACCESSION No. VOUCHER No COUNTRY
Thelephora americana AY219838.1 UAMH 9578 Canada
Thelephora anthocephala DQ974771.1 src614 USA
AF272927.1 TAA165304 Estonia
Thelephora caryophyllea GU234018.1 075453 Netherlands
AJ889980.1 TL-6566 Denmark
EU326158.1 105 Austria
EF655705.1 IB60087 Austria
Thelephora cf. penicillata EU819494.1 NAMA503 USA
Thelephora ganbajun EU696946.1 Gb279 China
Thelephora iqbalii JX241471 MH810 Pakistan
Thelephora palmata EU819443.1 JMP0085 USA
AF272919.1 TAA149550 Estonia
Thelephora aff. palmata AB509755.1 350-421 Japan
Thelephora penicillata U83484.1 LTT8, Bruns Herbarium, UCB USA
Thelephora pseudoterrestris AF272907.1 TAA159625 Estonia
AB453027.1 CU: Micro: Nan-M11 Thailand
Thelephora regularis U83485.1 OSC: JMT17371 USA
Thelephora sublilacina EF493288.1 UP161 Sweden
Thelephora terrestris HM189959.1 BB23-305-Ah-Pi-150506 (DNA64) Germany
HM189958.1 BB64-301-Oh-Pi-150506 (DNA43) Germany
Thelephoraceae sp. “Taylor #4 U83468.1 — USA
Thelephoroid sp. nw1 AF323110.1 — USA
Bold indicates sequence generated in this study; all others downloaded from GenBank.
Thelephora igbalii sp. nov. (Pakistan) ... 945
Collection and morphological analysis
During exploration of Himalayan moist temperate forests, an unidentified
Thelephora specimen associated with Abies pindrow was collected near a helipad,
in Khanspur, Abbotabad district, Khyber Pakhtunkhwa province, Pakistan. The
fresh specimen was morphologically characterized following Corner (1968), and
small portions of fresh hymenium were placed into 2% CTAB buffer and stored
at -20 °C for future molecular study. The remaining material was dried with a fan
heater overnight before storing in Ziploc bags. Basidiospores, basidia, basidioles
and sterigmata were observed in 1% KOH under a compound microscope and
measured using an ocular micrometer; drawings were made with the aid of a
camera Lucida. Dried specimens are deposited at the Herbarium of the Department
of Botany, University of the Punjab, Lahore (LAH).
Molecular analysis
Genomic DNA was extracted with a CTAB method (Gardes & Bruns 1993)
modified for silica emulsion binding and purified (Gene-Clean; Q-Biogene).
Polymerase chain reactions (PCR) for the ITS-nrDNA region were carried out
using the fungus-specific primers ITS1F/ITS4 following protocols described by
Gardes & Bruns (1993). PCR products were purified with QIAquick (Qiagen Inc.),
and sequenced by Macrogen (South Korea). Maximum Likelihood and Maximum
Parsimony methods generated a phylogeny comparing Thelephora iqbalii with other
thelephoroid species. A total of 22 sequences (21 downloaded from GenBank) were
used in the phylogenetic analyses (TABLE 1), and cladograms were recovered using
Mega5. Sequences were aligned with BioEdit version 7.0 (Hall 1999) and ClustalX
version 2.0 (Larkin et al. 2007) and corrected manually. All the gaps were treated
as ‘missing data. The ITS sequence generated from the holotype was deposited in
GenBank.
Results
Thelephora iqbalii Khalid & Hanif, sp. nov. FIG. 1
MycoBank MB 800830
Differs from Thelephora fucoides by its angular lobate basidiospores and from
T. penicillata by its coralloid basidiomata and its smaller basidiospores.
Type: Pakistan, Khyber Pakhtunkhwa province, Abbotabad district, Khanaspur, near
helipad, 1972 m a.s.l., associated with Abies pindrow (Royle ex D. Don) Royle, solitary
to gregarious, 22.VII.2010, M. Hanif, MH810 (Holotype, LAH 20810.2; GenBank
JX241471).
ErymMo.oey: dedicated to Prof. Dr. S.H. Iqbal in honor of his contributions to the
knowledge of fungi.
BASIDIOMA 4-5 x 2-4 cm, coralloid, individual branches 1-2 mm thick,
long narrow ridges, penicillate with fine hairs on long pointed tips, tip
946 ... Khalid & Hanif
—
Fic. 1. Thelephora iqbalii sp. nov. A. Basidiocarp in situ; B. Basidiospores; C. Basidia;
D. Hymenial tissue; E. Basidioles; F. Hair-like appendages of pointed tips; G. Tramal hyphae.
Bars: A = 1 cm, B—G = 10 um.
Thelephora iqbalii sp. nov. (Pakistan) ... 947
fibers typically serrate; branches pinkish white at the tip to grayish brown at
the base. CONTEXT solid, dry.
HyYPHAL SYSTEM monomitic, multi-septate, connective hyphae 6 um
diam., skeletal hyphae 5 um diam., tramal hyphae encrusted with globose
to sub-globose crystals, cyanescent in 1% KOH. Basip1a 50-75 x 8-10 um,
cylindrical to clavate, with basal clamps, smooth, 2-4 sterigmate, hyaline,
guttulate, light blue in KOH. BasipIoLes 34-40 x 4-8 um, smooth, thin
walled, hyaline, clavate, apex obtuse, light blue in KOH. BAsrp1osPoREs
6-8.5 x 5-6.5 um, reniform, angular lobate or scarcely lobate, thick-walled
and verrucose, tuberculate ornamentation, brown in lactic acid, light blue
in KOH.
ADDITIONAL MATERIAL EXAMINED Thelephora fucoides: MALAYSIA: PAHANG,
Cameron Highlands, 06.08.1934, E.J.H. Corner (E 00192884, holotype).
Phylogenetic analysis
Thelephora_caryophyllea_AJ889980.1
Thelephora_caryophyllea_EF655705.1
35 Thelephora_caryophyllea_EU326158.1
gg} 89 Thelephora_caryophyllea_GU234018.1
62 94 Thelephora_anthocephala_DQ974771.1 terminal clade 1
96 Thelephora_anthocephala_AF272927.1
Thelephoroid_sp._nw1_AF323110.1
60 @ Thelephora_iqbalii
65 Thelephora_penicillata_U83484.1
Thelephora_regularis_U83485.1 erases
86 Thelephora_palmata_EU819443.1 terminal clade 2
86 Thelephora_palmata_AF272919.1_
Thelephora_cf._penicillata_EU819494.1
= 59 51 TRS OR Paneer ese gw aT aN! ge a
53 79 Thelephora_pseudoterrestris_AB453027.1
65) Thelephora_ganbajun_EU696946.1
Thelephora_aff._palmata_AB509755.1
Thelephoraceae_sp._Taylor_#4°_U83468.1
74 Thelephora_terrestris_HM189959.1
84 Thelephora_americana_AY219838.1 | Clade Il
Thelephora_terrestris_HM189958.1
Thelephora_sublilacina_EF493288.1 | Outgroup
Fic. 2. Cladogram representing phylogenetic analysis inferred from Maximum Parsimony
(bootstrap percentages above the lines) and Maximum Likelihood (bootstrap percentages
below the lines).
Amplification of the ITS-nrDNA region produced one sequence of 654
bp. Initial BLAST analysis showed a 91% maximum identity with Thelephora
penicillata (GenBank U83484) with 100% query coverage.
948 ... Khalid & Hanif
Percent Identity
Thelephora_pseudoterrestris_AF272907
Thelephora_regularis_U83485
Thelephora_pseudoterrestris_AB453027
Thelephora_ganbajun_EU696946
Thelephora_palmata_EU819443
Thelephora_palmata_AF272919
Thelephora_cf_penicillata_EU819494
Thelephora_iqbalii
Thelephora_aff.__palmata_AB509755
Thelephora_americana_AY219838
Thelephora_sublilacina_EF 493288
Thelephora_terrestris_HM189959
Thelephora_terrestris_HM189958
Thelephora_penicillata_U83484
Thelephoraceae_sp._Taylor_#4'_U83468
Thelephoroid_sp._nw1_AF323110
Thelephora_caryophyllea_GU234018
Thelephora_caryophyllea_AJ889980
Thelephora_caryophyllea_EU326158
Thelephora_caryophyllea_EF655705
Thelephora_anthocephala_DQ974771
Thelephora_anthocephala_AF272927
Divergence
Fic. 3. Percent genetic divergence and percent similarity for Thelephora spp.
After trimming 84 unaligned characters from the 3’ and 5’ ends of the
789-bp ITS region, 705 characters were used for final analysis, of which
360 were conserved, 301 were variable but parsimony uninformative,
172 were parsimony informative, and 133 were singletons. The phylogenetic
tree generated two major clades (Fic. 2). Thelephora iqbalii clustered with
T. penicillata within terminal clade I of clade I, and T. anthocephala and
T: caryophyllea appeared as sister species to T! iqbalii. Thelephora iqbalii and
T. penicillata shared 91.8% genetic characters/nucleotides and showed an
8.1% genetic divergence (Fic. 3). However, we should note that the clustering
of all these species in terminal clade I of clade I (Fic. 2) was not supported
by both analyses.
Discussion
Eight Thelephora species have been previously recorded from Pakistan
(Ahmad 1956, 1972). Our new species, T. igbalii, which is supported
both by morphological and ITS sequence analysis, shares the basic
morphological characters with other Thelephora species: spiny to warty
and often lobed basidiospores, drab basidiocarp coloration, and blue to
green KOH reactions (Corner 1968). Hanif (2012) reported a rich diversity
of Thelephorales in subterranean ectomycorrhizal communities in HMT
forests, and T: iqbalii is expected to exhibit a similar ectomycorrhizal
association typical of Thelephora species elsewhere in Pakistan.
In Ahmad’s (1972) key to Thelephora species in Pakistan, T: iqbalii
would fall close to T: fucoides, which differs by its echinulate basidiospore
ornamentation (Corner 1968). Morphological and phylogenetic analyses
Thelephora iqbalii sp. nov. (Pakistan) ... 949
imply a close alliance between T. penicillata and T. igqbalii. However
T. penicillata can be distinguished by its resupinate basidiomes with
prostrate to suberect branches, its smaller basidiomata with margins that
are cristate, penicillate ramose-spiculose, or fimbriate, and its slightly larger
basidiospores (Corner 1968).
Acknowledgements
We are sincerely thankful to Prof. Juliano M. Baltazar (Centro de Ciéncias da
Natureza, Campus Lagoa do Sino, Universidade Federal de Sao Carlos (UFSCar), Buri,
Sao Paulo, Brazil) and Dr. Najam-ul-Sehar Afshan, (Centre for Undergraduate Studies,
University of the Punjab, Lahore, Pakistan) for acting as pre-submission reviewers.
We are also thankful for Dr. Stephan Helfer (Herbarium Royal Botanic Garden
Edinburgh, Scotland, UK) for sending the holotype of Thelephora fucoides for
morphological comparison.
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Ahmad S. 1956. Fungi of Pakistan, mon. I. Biological Society of Pakistan, Biological Laboratories,
Government College, Lahore, pp. 126.
Ahmad S. 1972. Basidiomycetes of Pakistan. Biological Society of Pakistan. Monograph 6. pp, 141.
Corner EJH. 1968. A monograph of Thelephora (Basidiomycetes). Beihefte zur Nova Hedwigia
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EI Karkouri K, Martin K Douzery E, Mousain D. 2005. Diversity of ectomycorrhizal
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persistence in nursery and plantations. FEMS Microbiology & Ecology 55: 146-158.
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Kirk PM, Ainsworth GC, Cannon PF, Minter DW. 2008. Ainsworth & Bisby’s Dictionary of fungi.
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Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H, Valentin F,
Wallace IM, Wilm A, Lopez R, Thompson JD, Gibson TJ, Higgins DG. 2007. Clustal W and
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Maas Geesteranus RA. 1971. Hydnaceous fungi of the eastern old world. Verh. K. Ned. Akad. Wet.
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Maas Geesteranus RA. 1975. Die terrestrichen Stachelpilze Europas. Verh. K. Ned. Akad. Wet. II
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Mittermeier RA, Turner WR, Larsen FW, Brooks TM, Gascon C. 2011. Global biodiversity
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Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J. 2000. Biodiversity hotspots
for conservation priorities. Nature403: 853-858.
Ramirez-Lopez I, Villegas-Rids M, Salas-Lizana R, Garibay-Orijel R, Alvarez-Manjarrez J. 2017.
Thelephora versatilis and Thelephora pseudoversatilis: two new cryptic species with polymorphic
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MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 951-955
https://doi.org/10.5248/132.951
Three new combinations in Drepanopeziza
for species on poplar
Amy Y. RossMAN’, W. CAVAN ALLEN’,
Lisa A. CASTLEBURY* & GERARD VERKLEY?
"Department of Botany & Plant Pathology, Oregon State University,
Corvallis, Oregon 97331, USA
*Mycology & Nematology Genetic Diversity & Biology Laboratory, USDA-ARS,
Beltsville, Maryland 20705, U.S.A.
>Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands
* CORRESPONDENCE TO: amydianer@yahoo.com
ABSTRACT— Three species of Drepanopeziza that cause diseases of poplars have been known
under different scientific names for their sexual and asexual morphs, which is no longer
allowed with the change to one scientific name for fungi. For each of these species, the oldest
epithet is provided by the asexual morph; however, neither of the generic names available
for the asexual morphs can be used for these species. As a consequence new combinations
for the three species are required and can be accomplished by applying the oldest asexual
morph epithet in the genus Drepanopeziza. The following new combinations are made:
Drepanopeziza brunnea, D. castagnei, and D. populi.
Key worps—fungal nomenclature, plant pathogen, pleomorphic fungi
Introduction
Following a careful study of type and other specimens, three species of
Drepanopeziza that cause diseases of poplar were delineated by Spiers &
Hopcroft (1998), who cited separate scientific names for their sexual and
asexual morphs. Because the International Code of Nomenclature for
algae, fungi and plants (ICN, McNeill et al. 2012) requires the adoption of
one scientific name for different morphs of one fungal taxon, these sexual
and asexual names now compete for use. For these three Drepanopeziza
952 ... Rossman & al.
species—D. populi-albae, D. populorum, and D. tremulae—the asexual morphs
provide the oldest epithets. Pirozynski (1974a,b,c) presented an account of
each of these species: D. tremulae as Marssonina brunnea, D. populi-albae as
Marssonina castagnei, and D. populorum as Marssonina populi, while Spiers
(1988) included a conidia-based key to these Marssonina species. Gremmen
(1965) recognized the same three taxa in Drepanopeziza but described one of
them as a new species, D. punctiformis Gremmen 1965, apparently unaware
of the earlier synonym D. tremulae Rimpau 1962.
In determining the generic names to use for competing synonyms, Rossman
et al. (2016) recommended that Drepanopeziza (Kleb.) Jaap 1914 be used rather
than the competing generic names Gloeosporidiella Petr. 1921 or Gloeosporium
Desm. & Mont. 1849. All three of these species of Drepanopeziza on poplar
have previously been placed in the genus Marssonina; however, these species
are not congeneric with the Marssonina type species, M. potentillae (Desm.)
Magnus 1906, now recognized as Diplocarpon earlianum (Ellis & Everh.)
EA. Wolf 1924 (Johnston et al. 2014). Diplocarpon EA. Wolf 1912 is now
regarded as the correct generic name for species formerly classified in
Marssonina Magnus 1906. Some authors use the Drepanopeziza names for
these species, as determined by Spiers & Hopcroft (1998); however, the asexual
morph names provide older epithets and are used more commonly (albeit in
the asexual morph genera that are no longer available for use). Rather than
conserve the less commonly used sexual morph names, it seems least disruptive
to make new combinations in Drepanopeziza using the asexual morph names,
as proposed here.
Taxonomy
Drepanopeziza brunnea (Ellis & Everh.) Rossman & W.C. Allen, comb. nov.
MycoBank MB822026
= Gloeosporium brunneum Ellis & Everh., J. Mycol. 5: 154 (1889).
Tye: U.S.A.: New Jersey, Newfield, on leaves of Populus candicans,
Aug 1889, Ellis & Everhart in North American Fungi, second ser. 2444
(Lectotype designated here, BPI 402970, under Marssonina brunnea).
= Marssonina brunnea (Ellis & Everh.) Magnus, Hedwigia 45: 88 (1906).
= Drepanopeziza tremulae Rimpau, Phytopathol. Z. 43: 288. (1962).
Type: Switzerland, Kt. Ziirich, am Uetliberg, bei Station Uitikon-Waldegg,
on Populus tremula, 7 Mai 1961, Rimpau (holotype, ZT-Myc 57794).
= Drepanopeziza punctiformis Gremmen, Nova Hedwigia 9: 172 (1965).
Type: Batava [the Netherlands], Meppel, in foliis Populi euroamericana
‘Serotina, 9.X.1961, Gremmen (holotype, herb. Gremmen 1811).
Drepanopeziza combs. nov. on poplar ... 953
Drepanopeziza brunnea causes a leaf spot disease of Populus that occurs
worldwide in temperate regions (Callan 1998, Farr & Rossman 2017). Spiers
& Hopcroft (1998) concluded that D. tremulae provided the oldest name for
the sexual morph of which D. punctiformis is a synonym. Pirozynski (1974a)
presented a description of this species using the asexual morph name Marssonina
brunnea based on Gloeosporium brunneum. Because neither Gloeosporium nor
Marssonina can be used as a generic name for this species, the name D. brunnea
is established based on Gloeosporium brunneum.
The type of G. brunneum was issued as Ellis & Everhart, North America Fungi
second series 2444, and the specimen at BPI is designated here as the lectotype.
According to Spiers & Hopcroft (1998), “type material of D. tremulae was not
deposited by Rimpau with other specimens at ZT and therefore was probably
lost” However, the holotype specimen was later found at ZT (R. Berndt pers.
comm. 31 Jan 2017).
Drepanopeziza castagnei (Desm. & Mont.) Rossman & W.C. Allen, comb. nov.
MycoBank MB822027
= Gloeosporium castagnei Desm. & Mont., Ann. Sci. Nat. Bot., sér. 3, 12: 295 (1849).
TyPE: France, Aix, on Populus alba, 1852, Castagne, ex Herb. Bois, Shear Type
and Rarities Ser. 1 (Lectotype designated here, BPI barcode 797921).
= Marssonina castagnei (Desm. & Mont.) Magnus, Hedwigia 45: 88 (1906).
= Pseudopeziza populi-albae Kleb., Haupt- und Nebenfruchtform. Ascomyz. 1:
344 (1920) [“1918”]. Type: Germany, Berlin, Dahlem, Garten der Biologischen
Reichsanstalt, on leaves of Populus alba, Oct 1913, Laubert (holotype, HBG).
= Drepanopeziza populi-albae (Kleb.) Nannf., Nova Acta R.
Soc. Scient. Upsal., Ser. 4 8(2): 170 (1932).
Drepanopeziza castagnei causes a leaf disease of poplar that is widespread in
temperate regions (Callan 1998, Farr & Rossman 2017). Spiers & Hopcroft
(1998) considered Marssonina castagnei based on Gloeosporium castagnei to be
the asexual morph of D. populi-albae. Pirozynski (1974b) provided an account
of this species under the asexual morph name Marssonina castagnei. Because
neither Gloeosporium nor Marssonina can be used as a generic name for this
species, it is least disruptive to make a new combination in Drepanopeziza with
the oldest and familiar epithet castagnei.
Although Spiers (1988) mentioned that “Type and isotype material of
M. castagnei was obtained from...” BR and PC, he did not present details
of the specimens, so his work cannot be considered lectotypification of
the name G. castagnei. A specimen matching the protologue exists at BPI,
and this specimen is designated here as the lectotype. According to Stafleu
& Cowan (1976) Klebahn’s specimens are housed at B, but Robert Licking
954 ... Rossman & al.
(pers. comm.) reported that the type of Pseudopeziza populi-albae is not
found there. Rather, a specimen matching the protologue was located at
HBG.
Drepanopeziza populi (Lib.) Rossman & W.C. Allen, comb. nov.
MycoBAnk MB 822028
= Leptothyrium populi Lib., Pl. Crypt. Arduenna, fasc. 3: no. 257 (1834).
TyPE: France, on Populus, autumn (Lectotype designated here, FH).
= Marssonina populi (Lib.) Magnus, Hedwigia 45: 88 (1906).
= Trochila populorum Desm., Pl. Crypt. Nord France Ed. (2) 2, no. 451 (21 Mar 1857).
TYPE: France, on dried leaves of Populus nigra, spring, 1857, Grance,
Herb. Desmaziéres (lectotype, PC; isolectotypes, BR, BPI-bound).
= Drepanopeziza populorum (Desm.) Hohn., Ann. Mycol. 15: 332 (1917).
Drepanopeziza populi causes a leaf disease of poplar that is widespread in
temperate regions (Callan 1998, Farr & Rossman 2017). Spiers & Hopcroft
(1998) considered Marssonina populi, based on Leptothyrium populi, to be the
asexual morph of D. populorum. This fungus was described and illustrated
by Pirozynski (1974c) using the asexual morph name Marssonina populi. As
neither Gloeosporium nor Marssonina is available as a generic name for this
species, it is least disruptive to combine the well-used epithet in the appropriate
genus Drepanopeziza as D. popuili.
The name Leptothyrium populi was published in an exsiccati number that
includes a description; the title page of the fascicle carries the date 1834 (Pfister
1985) but Sayre (1969) dated it as 1837 on the basis of reviews. The type
specimen of Trochila populorum was issued as part of Desmaziéres’ exsiccati on
21 March 1857, according to Pfister (1985). This number includes a description
on the label and thus should be regarded as the original description and type
specimen. This name was also included in an article published later in March
1858 in Desmaziéres (1858), which has mistakenly been regarded as the place
of publication. A lectotype was designated by Spiers & Hopcroft (1998).
Acknowledgements
We thank curatorial staff at several herbaria for their assistance, specifically Reinhard
Berndt (ZT), Robert Licking (B), Donald Pfister (FH), and Matthias Schultz (HBG). In
addition we appreciate the thoughtful reviews of Jeffrey Stone (Oregon State University,
Corvallis) and Peter Johnston (Landcare Research, Auckland, New Zealand).
Mention of trade names or commercial products in this publication is solely for
the purpose of providing specific information and does not imply recommendation
or endorsement by the United States Department of Agriculture. USDA is an equal
opportunity provider and employer.
Drepanopeziza combs. nov. on poplar ... 955
Literature cited
Callan BE. 1998. Diseases of Populus in British Columbia: a diagnostic manual. Natural Resources
Canada, Canadian Forest Service.
Desmaziéres JBHJ. 1858 [“1857”]. Vingt-quatriéme notice sur les plantes cryptogames
récemment découvertes (suite). Bulletin de la Société Botanique de France 4(9): 858-863.
https://doi.org/10.1080/00378941.1857.10829049
Farr DE, Rossman AY. 2017. U.S. National Fungus Collections Databases, retrieved 20 Jan 2017,
from https://nt.ars-grin.gov/fungaldatabases
Gremmen J. 1965. Three poplar-inhabiting Drepanopeziza species and their life-history. Nova
Hedwigia 9: 170-176.
Johnston PR, Seifert KA, Stone JK, Rossman AY, Marvanova L. 2014. Recommendations on
generic names competing for use in Leotiomycetes (Ascomycota). IMA Fungus 5: 91-120.
https://doi.org/10.5598/imafungus.2014.05.01.11
McNeill J, Barrie FE Buck WR, Demoulin V, Greuter W, et al. (eds). 2012. International Code of
Nomenclature for algae, fungi, and plants (Melbourne Code). [Regnum Vegetabile no. 154.]
K6nigstein: Koeltz Scientific Books.
Pirozynski KA. 1974a. Marssonina brunnea. Fungi Canadenses 13. 2 p.
Pirozynski KA. 1974b. Marssonina castagnei. Fungi Canadenses 14. 2 p.
Pirozynski KA. 1974c. Marssonina populi. Fungi Canadenses 15. 2 p.
Pfister DH. 1985. A bibliographic account of exsiccatae containing fungi. Mycotaxon 23: 1-139.
Rossman AY, Allen WC, Braun U, Castlebury LA, Chaverri P, et al. 2016. Overlooked competing
asexual and sexually typified generic names of Ascomycota with recommendations for their
use or protection. IMA Fungus 7: 289-308. https://doi.org/10.5598/imafungus.2016.07.02.09
Sayre G. 1969. Cryptogamae exsiccatae--an annotated bibliography of published exsiccatae of
algae, lichenes, hepaticae, and musci. Memoirs of the New York Botanical Garden 19. 174 p.
Spiers AG. 1988. Comparative studies of type and herbarium specimens of Marssonina
species pathogenic to poplars. European Journal of Forest Pathology 18: 140-156.
https://doi.org/10.1111/j.1439-0329.1988.tb00912.x
Spiers AG, Hopcroft DH. 1998. Morphology of Drepanopeziza species pathogenic to poplars.
Mycological Research 102: 1025-1037. https://doi.org/10.1017/S0953756297005972
Stafleu FA, Cowan RS. 1976. Taxonomic literature. A selective guide to botanical publications and
collections with dates, commentaries and types. Vol. 1. Bohn, Scheltema and Holkema.
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 957-969
https://doi.org/10.5248/132.957
Key to the lichen families Pyrenulaceae and Trypetheliaceae in
Vietnam, with eight new records
SANTOSH JOSHI’, D. K. UPRETI’ & JAE-SEOUN HuR**
'Lichenology Laboratory, CSIR-National Botanical Research Institute,
Rana Pratap Marg, Lucknow (UP)-226001, India
?Korean Lichen Research Institute, Sunchon National University, Suncheon-540 950, Korea
* CORRESPONDENCE TO: jshur1@sunchon.ac.kr
AssTRACT—An identification key is presented for the members of the lichen families
Pyrenulaceae and Trypetheliaceae in Vietnam. Eight pyrenocarpous species (in
Anthracothecium, Astrothelium, Lithothelium, and Pyrenula) collected from Nam Cat Tien
National Park, are new records for Vietnam. Taxonomic characters of the species are given
along with ecology, distribution, and illustrations.
Key worps—Bathelium, Nigrovothelium, taxonomy, Trypethelium
Introduction
Pyrenocarpous lichens in the tropics belong mainly to the families
Porinaceae, Pyrenulaceae, Strigulaceae, and Trypetheliaceae and are among the
major constituents of epiphytic lichen communities in primary rain forests
(Aptroot 2009). Although little remains of Vietnam's primary forests, the
protected areas, reserve forests, and national parks still conserve significant
stretches of old-growth rain forests, particularly in the southern part of the
country. The present inventory is the result of ongoing research conducted in
Nam Cat Tien National Park situated in southern Vietnam and provides new
records of eight pyrenocarpous lichens in Trypetheliaceae and Pyrenulaceae for
Vietnam. Both families have been the focus of increasing global interest in recent
years. Currently, 234 species in 11 genera are accepted in Pyrenulaceae, and 418
species in 15 genera in Trypetheliaceae (Aptroot 2012, Aptroot & Licking 2016,
Licking et al. 2016). From Vietnam 24 species of Pyrenulaceae have been
958 ... Joshi, Upreti & Hur
recorded: Anthracothecium macrosporum (Hepp) Mill. Arg., A. prasinum
(Eschw.) R.C. Harris; Pyrenula aggregata (Fée) Fée, P. anomala (Ach.) Vain.,
P. aspistea (Ach.) Ach., P. balia (Kremp.) R.C. Harris, P breutelii (Mull. Arg.)
Aptroot, P brunnea Fée, P. duplicans (Nyl.) Aptroot, P fetivica (Kremp.) Mill.
Arg., P. mamillana (Ach.) Trevis., P. massariospora (Starback) R.C. Harris,
P. microcarpa Mull. Arg., P. nitidella (Schaer.) Mull. Arg., P nitidula (Bres.)
R.C. Harris, P. ochraceoflava (Nyl.) R.C. Harris, P. parvinuclea (Meyen &
Flot.) Aptroot, P pyrenuloides (Mont.) R.C. Harris, P quassiicola (Fée) Fée,
P. scutata (Stirt.) Zahlbr., P sexlocularis (Nyl.) Mill. Arg., P subglabrata
(Nyl.) Mull. Arg., P sublaevigata (Patw. & Makhija) Upreti, P thelemorpha
Tuck.—while 16 species have been recorded in Trypetheliaceae: Astrothelium
aeneum (Eschw.) Aptroot & Liicking, A. cinnamomeum (Eschw.) Mull. Arg.,
A. clypeatum Aptroot & Gueidan, A. megaspermum (Mont.) Aptroot & Licking,
A. nitidiusculum (Nyl.) Aptroot & Licking, A. phlyctaena (Fée) Aptroot &
Licking [= Trypethelium ochroleucum (Eschw.) Nyl.], A. porosum (Ach.)
Aptroot & Liicking, A. variolosum (Ach.) Mull. Arg.; Bathelium albidoporum
(Makhija & Patw.) R.C. Harris, B. lineare (C.W. Dodge) R.C. Harris,
B. porinosporum Licking et al.; Nigrovothelium tropicum (Ach.) Licking et al.;
Trypethelium eluteriae Spreng., T. epileucodes Nyl., T. infraeluteriae Aptroot
& Gueidan, T. krempelhuberi Makhija & Patw. (Aptroot & Sparrius 2006,
Aptroot 2012, Aptroot et al. 2016, Liicking et al. 2016).
In addition to some previously reported species, we examined some
interesting pyrenocarpous lichens that we report here as new Vietnamese
records: Anthracothecium interlatens, Astrothelium galligenum,
A. inspersogalbineum, A. subaequans, Lithothelium obtectum, Pyrenula
circumfiniens, P. laetior, and P. mastophora. An artificial key to all known
species in Vietnam representing Pyrenulaceae and Trypetheliaceae is also
presented.
Materials & methods
The Vietnamese material was collected by Dr. J.-S. Hur and J.J. Woo in December
2015 from the lowland areas in southern Vietnam having warm and humid preserved
forests with a sub-oceanic climate. The material was deposited in the lichen herbarium
of the Korean Lichen Research Institute, Suncheon, South Korea (KoLRI), and studied
in the Lichenology laboratory of CSIR-National Botanical Research Institute, Lucknow,
India. The morphological characters were identified and measured using a MSZ-TR
dissecting microscope and a Leica DM 500 compound microscope. Standard protocols
were followed to analyse chemistry (Orange et al. 2010). Amyloidity (I+ or I-) of
internal structures was tested using Lugol's solution. Images were taken using a Zeiss
Axiocam ERc5s and Al(AX10) microscope. The key includes some characters that
Pyrenulaceae & Trypetheliaceae in Vietnam ... 959
were not apparent in our specimens but were derived mainly from Upreti (1991, 1998),
Aptroot et al. (2008), Aptroot (2012), and Aptroot & Liicking (2016).
Taxonomy
Anthracothecium interlatens (Nyl.) Aptroot, Lichenologist 44: 35, 2012. PL. 1A
Thallus epi- to endoperidermal, pale green to fawn, corticate, continuous,
matt to slightly glossy, reflecting bark texture, delimited by blackish prothallus,
100-120 um thick; cortex 50-60 um thick; photobiont trentepohlioid, layer
distinct, <50 um thick; medulla not apparent; perithecia semi-immersed to
emergent, black, scattered to aggregate with shared ostiole, 0.5-0.7 mm in
diam.; perithecial wall complete, carbonized, 45-80 um thick; ostiole apical,
0.071-0.1 mm in diam; hamathecium hyaline, clear, with unbranched filaments;
asci 2-spored, 200-230 x 45-52 um, I-; ascospores mostly hyaline to grayish at
maturity, oblong to slightly curved, muriform, 100-132 x 32-42 um, I-.
CHEMISTRY—No lichen substances present.
DISTRIBUTION & ECOLOGY—Pantropical (Aptroot 2012). In Vietnam, the
species was found growing in association with Ocellularia spp. in irregular
patches on uneven bark surfaces.
SPECIMEN EXAMINED: VIETNAM. DonG Nat PRovINCcE: Tan Phu district, Cat
Tien National Park, forest red soil area, 11°24’20”N 107°17'19’E, alt. 256 m, on bark,
17 December 2015, Hur & Woo VN150006 (KoLRI).
REMARKS—Anthracothecium gregale (C. Knight) Aptroot is morphologically
close but produces 6-8-spored asci (Aptroot 2012). Only a small amount of
A. interlatens was collected in only a small amount from the national park, but
the Vietnamese specimen was easily identifiable.
Astrothelium galligenum (Aptroot) Aptroot & Licking,
Lichenologist 48: 861, 2016. PL. 1B
Thallus epiperidermal, pale greenish gray to greenish gray, smooth,
continuous, glossy, delimited by blackish prothallus, <180 um thick; cortex
distinct, continuous, 50-60 um thick; photobiont trentepohlioid, layer distinct,
continuous, 40-50 um thick; medulla white crystalline, reaching <70 um;
ascomata immersed to slightly emergent, trypethelioid, scattered, solitary to
slightly confluent, 0.2-0.3 mm in diam., pseudostromatic; pseudostromata
off-white, irregular in shape; ostiole apical, black, 0.01-0.03 mm in diam.;
perithecial wall complete, carbonized, 90-100 um thick hamathecium hyaline,
inspersed with oil droplets, with anastomosing filaments; asci 8-spored, 140-
170 x 20-30 um, I-; ascospores uni- to biseriate, fusiform, hyaline, transversely
7-8-septate with diamond shaped lumina, 40-47 x 10-12 um, I-.
960 ... Joshi, Upreti & Hur
CHEMISTRY— thallus and ascomata UV-, hamathecium oil droplets
K+ purple; anthraquinone present.
DISTRIBUTION & ECOLOGY—Eastern Palaeotropics (Aptroot & Liicking
2016). This species is apparently rare in the national park, where it was
growing in association with Bathelium mastoideum and Sarcographa spp.
on trees with somewhat undulating thick smooth bark.
SPECIMEN EXAMINED: VIETNAM. Dong Nal PROVINCE: Tan Phu district, Nam Cat
Tien National Park, in forest red soil area, 11°23’49”N 107°17’38’E, alt. 206 m, on bark,
17 December 2015, Hur & Woo VN150125 (KoLRI).
REMARKS—Astrothelium ubianense (Vain.) Aptroot & Licking is similar,
but has a clear hamathecium and lacks anthraquinone (Aptroot & Licking
2016).
Astrothelium inspersogalbineum Aptroot & Weerakoon,
Lichenologist 48: 618, 2016. PIE AC
Thallus epiperidermal, greenish gray to yellowish green, smooth,
continuous, glossy, delimited by blackish prothallus, 100-150 um thick;
cortex distinct, 30-40 um thick photobiont trentepohlioid, layer distinct,
continuous, 50-60 um thick; medulla endoperidermal; ascomata emergent,
trypethelioid, scattered, solitary to slightly aggregate then open by a
common ostiole, mostly covered by thallus, 0.6-0.7 mm in diam.; ostiole
apical to slightly eccentric, black, 0.08-0.1 mm in diam.; perithecial wall
complete, carbonized, 100-150 um thick laterally, reaching <350 um basally;
hamathecium hyaline, inspersed with oil droplets, with anastomosing
filaments, placed deep into the ascomata; asci 8-spored, 40-50 x 10-12 um,
I-; ascospores mostly uniseriate, fusiform, hyaline, transversely 3-septate
with diamond shaped lumina, 20-22 x 7-9 um, I-.
CHEMISTRY—Thallus and ascomata UV+ yellow, K+ _ purple;
lichexanthone and anthraquinone present.
DISTRIBUTION & ECOLOGY—This is only the second report of this taxon,
originally described from Singapore (Aptroot et al. 2016). In Vietnam, it
was growing luxuriantly in association with Bathelium mastoideum on
thick smooth-barked trees.
SPECIMEN EXAMINED: VIETNAM. Dona Nal PROVINCE: Tan Phu district, Nam Cat
Tien National Park, in forest red soil area, 11°23’36”N 107°17'56”E, alt. 181 m, on bark,
17 December 2015, Hur & Woo VN150126 (KoLRI).
REMARKS— This species is close to Astrothelium macrocarpum (Fée) Aptroot
& Liicking, which differs in its clear hymenium and pseudostromatic
ascomata (Aptroot 2012). Our Vietnamese A. inspersogalbineum specimen,
Pyrenulaceae & Trypetheliaceae in Vietnam ... 961
with a K+ purplish thallus, differs slightly from the K— holotype (Aptroot
& Liicking 2016).
Astrothelium subaequans Mill. Arg., Bot. Jahrb. Syst. 6: 383, 1885. PL. 1D
Thallus epiperidermal, olive green, bullate, uneven, hard, continuous, dull
to + glossy, delimited by blackish prothallus, 200-300 um thick, photobiont
trentepohlioid, layer distinct, continuous, 40-50 um thick; medulla white,
crystalline, 200-350 um thick; ascomata astrothelioid, scattered, solitary to
aggregate and pseudostromatic, covered by thallus, surrounded by yellow-
orange pigments, 0.7-1 mm in diam.; ostiole slightly eccentric, black,
0.5-0.1 mm in diam.; perithecial wall complete, carbonized, laterally
100-130 um thick and basally reaching <600 um, separated from the thallus
by a slit; hamathecium hyaline, clear with anastomosing filaments, placed
deep into the ascomata; asci 8-spored, 145-155 x 40-47 um, I-; ascospores
biseriate, oval, hyaline, muriform, 60-65 x 18-21 um, [-.
CHEMISTRY— Thallus UV-, ascomata UV+ reddish, pigment K+ purple;
anthraquinone present.
DISTRIBUTION & ECOLOGY—Previously known from the Neotropics
(Aptroot 2016). In Vietnam, the species was growing richly in large patches
on trees with thick and hard bark.
SPECIMEN EXAMINED: VIETNAM. Dong NaI PROVINCE: Tan Phu district, Nam Cat
Tien National Park, in forest red soil area, 11°24’20”N 107°17'19”E, alt. 256 m, on bark,
17 December 2015, Hur & Woo VN150054 (KoLRI).
REMARKS—This species is close to Astrothelium praetervisum (Mull. Arg.)
Aptroot & Liicking, which differs in producing pseudostromatic ascomata
(Aptroot & Licking 2016). Our Vietnamese A. subaequans specimen is
characterized by distinctly muriform ascospores that differ slightly from
the submuriform ascospores described by Aptroot & Liticking (2016).
Lithothelium obtectum (Mill. Arg.) Aptroot,
Biblioth. Lichenol. 44: 62, 1991. PL. 1E
Thallus epiperidermal, grayish green, corticate, continuous, dull to
slightly glossy, delimited by black prothallus, <100 um thick; cortex 20-30
um thick; photobiont trentepohlioid, layer 40-50 um thick; medulla not
apparent; perithecia emergent, black, irregularly in shape to shortly elongate,
numerous, solitary to aggregated in groups of 2-3 joint together with fused
ostiole, 0.2-0.7 mm long; ostiole eccentric to laterally placed, 0.05-0.1 mm in
diam.; perithecial wall complete, carbonized, 90-100 um thick; hamathecium
hyaline, clear, with unbranched filaments; asci 8-spored, 60-90 x 8-12 um,
962 ... Joshi, Upreti & Hur
I-; ascospores uniseriate, fusiform, hyaline, transversely 3-septate, 10-15 x
3-4 um, I-.
CHEMISTRY—No lichen substances present.
DISTRIBUTION & ECOLOGY—Pantropical (Aptroot et al. 2008). In Vietnam,
the species was collected from hard, rather rough-barked trees, where it was
growing luxuriantly with crustose lichen taxa, probably Graphidaceae.
SPECIMENS EXAMINED: VIETNAM. Dong NAI PROVINCE: Tan Phu district, Nam Cat
Tien National Park, near Crocodile lake, 11°27’25”N 107°21’4’E, alt. 156 m, on bark, 19
December 2015, Hur & Woo VN150400, VN150402 (KoLRI).
REMARKS— The examined specimen differs from the type of Sagedia obtecta
Mill. Arg. (especially in thallus formation and number of fused ascomata).
It is possible that the ecorticate thin thallus and solitary ascomata describing
the type are features taken from a damaged specimen with an eroded thallus.
Lithothelium decumbens (Mull. Arg.) Aptroot is close to L. obtectum but differs
in producing brown ascospores (Aptroot 2012).
Pyrenula circumfiniens Vain.,
Ann. Acad. Sci. Fenn., Ser. A, 6(7): 195, 1915. PL. 1F
Thallus epiperidermal, olive green to dark green, corticate, continuous,
glossy, delimited by black prothallus, 100-150 um thick; cortex 40-60 um
thick; photobiont trentepohlioid, layer 50-70 um thick; medulla not apparent;
perithecia semi-immersed, black, numerous, solitary to sometimes aggregated
in 2-3 groups, 0.3-0.5 mm in diam.; ostiole eccentric to laterally placed
in different directions, 0.01-0.2 mm in diam.; perithecial wall complete,
carbonized, 90-100 um thick; hamathecium hyaline, clear, with unbranched
filaments; asci 8-spored, 60-90 x 8-12 um, I-; ascospores uniseriate, fusiform,
gray brown, transversely 3-septate, terminal lumina directly against the
exospores wall, 13-18 x 4-6 um, I-.
CHEMISTRY—No lichen substances present.
DISTRIBUTION & ECOLOGY—Pantropical (Aptroot 2012). In Vietnam,
Pyrenula circumfiniens was flourishing well around trees having wide girth.
The species was spread largely in irregular patches and intermingled marginally
with other pyrenocarpous lichens.
SPECIMEN EXAMINED: VIETNAM. Dong Nal PROVINCE: Tan Phu district, Nam Cat
Tien National Park, near rock stream, 11°26’39”N 107°24’22’E, alt. 160 m on bark,
18 December 2015, Hur & Woo VN150250 (KoLRI).
REMARKS— The ascospore sizes in our sample are somewhat close to those in
Pyrenula elliptica Mill. Arg., which is distinguished by its slightly larger (15-18 x
6-8 um) ascospores and distribution restricted to the Neotropics (Aptroot 2012).
Pyrenulaceae & Trypetheliaceae in Vietnam ... 963
Pyrenula laetior Mill. Arg., Bot. Jahrb. Syst. 6: 413, 1885. PL. 1G
Thallus epi- to endoperidermal, brownish green to reddish brown, corticate,
continuous, + dull, delimited by black prothallus, <100 um thick; cortex 20-25
um thick; photobiont trentepohlioid, layer <30 um thick; medulla indistinct,
mostly endoperidermal; perithecia semi-emergent, black, numerous, simple
to aggregate, 0.2-0.3 mm in diam.; ostiole apical, 0.05-0.07 mm in diam.;
perithecial wall complete, carbonized, 50-80 um thick; hamathecium hyaline,
inspersed with oil droplets, with unbranched filaments; asci 8-spored,
40-60 x 8-6 um, I-; ascospores uniseriate, fusiform, gray brown, transversely
3-septate, terminal lumina separated from the exospore wall by endospore
thickening,13-15 x 4-6 um, I-.
CHEMISTRY—No lichen substances present.
DISTRIBUTION & ECOLOGY—Previously known from the Neotropics
(Aptroot 2012). In Vietnam, the species is found on thin papery bark flaking
away from tree trunks.
SPECIMEN EXAMINED: VIETNAM. DonG NaI PROVINCE: Tan Phu district, Cat Tien
National Park, forest red soil area, 11°24’23”N 107°17’40’E, alt. 288 m, on bark, 17
December 2015, Hur & Woo VN150083 (KoLRI).
ReMARKS—Pyrenula subglabrata, previously reported from Vietnam, is
morphologically similar to P laetior but can be distinguished by its longer
(18-20 um) ascospores (Aptroot 2012). Our material, which produces very
short (<15 um) ascospores fits well within the range of the new world Pyrenula
laetior.
Pyrenula mastophora (Nyl.) Mill. Arg., Flora 66: 426, 1883. PL. 1H
Thallus epiperidermal, green to grayish green, corticate, continuous,
smooth, delimited by a black prothallus, <150 um thick; cortex 35-40 um
thick; photobiont trentepohlioid, layer <45 um thick; medulla indistinct;
perithecia semi-immersed to semi-emergent, initially covered by thallus, black,
numerous, scattered, 0.5-0.8 mm in diam.; ostiole apical, indistinct to 0.5 mm
in diam.; perithecial wall complete, carbonized, 60-90 um thick; hamathecium
hyaline, clear, with unbranched filaments; asci 8-spored, 150-160 x 20-26 um,
I-; ascospores mostly uniseriate, fusiform, gray brown, transversely 3-septate,
terminal lumina separated from the exospore wall by endospore thickening,
28-32 x 11-13 um, I-.
CHEMISTRY—No lichen substances present.
DISTRIBUTION & ECOLOGY—Pantropical (Aptroot 2012). The Vietnamese
specimen broadly colonized on trees with thick smooth bark.
964 ... Joshi, Upreti & Hur
SPECIMEN EXAMINED: VIETNAM. Donec Nal PROVINCE: Tan Phu district,
Nam Cat Tien National Park, near rock stream, 11°26’35”N 107°24’19’E, alt.
150 m on bark, 18 December 2015, Hur & Woo VN150216 (KoLRI).
REMARKS—Pyrenula quassiicola is similar but is distinguished from
P. mastophora by its partly immersed perithecia and thallus producing
pseudocyphellae (Aptroot 2012).
Key to species in Pyrenulaceae and Trypetheliaceae
recorded from Vietnam
DUASCOSPOLES YAH Sle oleae eect ec teeh peste hor See i RPE gr REED er AES pr SESE dls Z
fh iXscosporespiemented .. . Mince: Siw «Mews 2 Wenis« Wows Whwioe VEcwnia ey Seema 2 peer Al 21
2, Ascospores. transversélyseptgtel. ose. 4c 52sec eeu y ene yas syne sy aae he aee ey ae 3
De ASCOSPOLESATUTIOBIN ter x15 (5p eect ee nate ete cee E). ieee eect Ep stce ekjestee Epes 19
3. Ascospore lumina rectangular to oval at maturity .....................0.0008. 4
3.Ascospore lumina diamond-shaped wij. Sele olie linea deg fis: siguat hy idea hos asd gs slide pee 10
4. Ascospores 3-septate, 10-15 x 3-4 um; ostiole lateral ...... Lithothelium obtectum
4, Ascospores:>3-septatexostiole:apical ... 2s .45 ees yin d ewe 2 ving 04 bina 0 Fe ee ee et 5
5, Ascomata solitary: to irregularly-confluent. «ix <4. 5 twas eee tee ee 6
Sv ASCOMAtA-WSCMUOSLLONIALIC Monat oh nal estan etc etna Aenea et A Med te 7
6. Ostiole usually white pruinose; ascospores transversely 5—7-septate,
Dis ASO GeO NTIS as Sa ays, sudled su Bhs epee peed a atasey dime es sea Bathelium albidoporum
6. Ostiole non-pruinose; ascospores transversely 3-5-septate,
ZOE BEI G ss (TDER gs saat a ak Fate Sad s sap So’ Ene asks geese cet Bathelium porinosporum
7. Pseudostromata lacking anthraquinone (all white with small dark ostiole spots;
ascospores 5-13-septate, 30-60 x 5-13 um) ........... Trypethelium epileucodes
AP seucostrotata witht antnnaquinvOnes 4825 ot, a's ek bts eae PE ah ed otek 8
8. Ostiolar area broad, disc shaped with margins producing
black papillae arranged in circular shapes
(ascospores 13-15-septate, 59-72 x 13-15 um) .... Trypethelium krempelhuberi
8. Ostiolat-area harrow, visibleas:small dots? ww o5 aisha sob a wh eek ee aot 9
9. Pseudostromata immersed to erumpent;
ascospores 7-9-septate, 37-42 x 9-llum......... Trypethelium infraeluteriae
9. Pseudostromata prominent to sessile;
ascospores 9-13-septate, 37-52 x 8-llum............ Trypethelium eluteriae
PLATE. 1. Specimens of pyrenocarpous lichens from Vietnam. A. Anthracothecium interlatens
(KoLRI VN150006); B. Astrothelium galligenum (KoLRI VN150125); C. A. inspersogalbineum
(KoLRI VN150126); D. A. subaequans (KoLRI VN150054); E. Lithothelium obtectum (KoLRI
VN150400); E Pyrenula circumfiniens (KoLRI VN150250); G. P. laetior (KoLRI VN150083);
H. P mastophora (KoLRI VN150216). Scale bars: 1 mm.
Pyrenulaceae & Trypetheliaceae in Vietnam ... 965
966 ... Joshi, Upreti & Hur
10.
10.
11.
ile
12
12.
13.
Ns 8
14.
14.
BS;
15.
16.
16.
| Ws
17.
18.
18.
19;
13:
20.
2
i=)
2
2
2
—_—_ —_—
N
2
N
Ascospores 7-8-septate (40-47 x 10-12 um) .......... Astrothelium galligenum
PRS COSPOTESOASCPLALE 25 2 screens acer eae te eto Pt Pe fs coh SEs tt AR 1!
Hamathecium inspersed with oil droplets ......... 0... cece ee eee eee eee 12
PianratHeeniin cleats 08 tes Ae henge sets A eRe Se RAL Re Rs vel RS lt ReS cits 14
. Ostiole lateral (ascospores 20-22 x 7-9 um;
anthraquinone and lichexanthone present) .... Astrothelium inspersogalbineum
CS TEOLE AACA nar ox sole Paget ap te Bek rs PG eng cae Pema Peer poe ee EER 13
Ascomata solitary; ascospores 17-27 x 7-10 um;
anthraquinone and lichexanthone both absent ........ Astrothelium clypeatum
Ascomata diffusely pseudostromatic; ascospores 17-27 x 7-10 um;
anthraquinone present, lichexanthone absent .......... Astrothelium porosum
Ostioleapicale, 450d hs nset eck sitet dst leads ap seg bos onus mbes a b-eed ables b-eleee | 15
MSO Le Patel Ae a. Acts eo. ete p-cetelee-tictetine- aeine nak cath gti gay cach on eae bg SOEs eg AOE 18
Ascomata distinctly pseudostromatic (ascospores 15-27 x 7-10 um;
anthraquinone absent, lichexanthone present) ....... Astrothelium phlyctaena
Ascomata solitary to irregularly confluent or diffusely pseudostromatic ....... 16
Ascomata solitary to diffusely pseudostromatic; anthraquinone present,
lichexanthone absent (ascospores 20-27 x 7-10 um)..... Astrothelium aeneum
Ascomata solitary to irregularly confluent or crowded; anthraquinone and
Hehe xanthone Doth -abSene 40-8 k eee tinh ake mn hod enw Balegenn Souk ea Roky Pade ete Bore 17
Ascomata sessile, fully exposed, barrel-shaped, black;
ascospores 20-25 x 7-10 UM... 0... eee ee eee eee Nigrovothelium tropicum
Ascomata erumpent with broad white area surrounding the ostiole
or entirely white; ascospores 15-27 x 7-10 um..... Astrothelium nitidiusculum
Ascospores 20-26 x 7-9 um;
lichexanthone present, anthraquinone absent ........ Astrothelium variolosum
Ascospores 23-30 x 6-10 um;
lichexanthone absent, anthraquinone present ..... Astrothelium cinnamomeum
Ascospore lumina rectangular to oval at maturity
(1OQ=12. ool SASTRY osx cce aih yest wite'y eee een oa nate alloy Bathelium lineare
AScospore luimmiinaiamand=shaped! 245 eenp5e 0 donceps8s-vuonst4he pany the eaggtdhins jqate ators yates 20
Ostiole apical; hamathecium inspersed with oil droplets; ascospores mostly 4/ascus,
140-220 x 30-75 um; anthraquinone absent ....... Astrothelium megaspermum
. Ostiole lateral; hamathecium clear; ascospores 8/ascus,
60-65 x 18-21 um; anthraquinone present .......... Astrothelium subaequans
. Ascospores transversely séptatew, .a5 + aces een eye k eoayee 4 elas ele 4 Sings oe lig 22
WING COSPOLE SIU TILOLITE wa. aak ete aceath et aca e te wee tD. cee ge fe ieee EPs ence. se Ee se ge Leste EE 39
. Ostiole lateral, pointing in different directions
(ascospores 3-septate, 13-18 x 4-6 um)............... Pyrenula circumfiniens
1B See) coe) 6 (cre Leis Si ees, ger CORR rig Rrra SI Gn, AAO, Ont UR A ORR, A rt TERE ADEA 23
23
23,
24.
24.
pep
25;
26.
26.
pi
2
28.
28.
29,
29
30.
30.
3
3)
—_ —
os
32.
Dos
AS}
34.
34.
35.
oa.
36.
36.
OL
ats
38.
38.
39,
39,
Pyrenulaceae & Trypetheliaceae in Vietnam ... 967
Ascospores 4-7-septate, with orange oil at maturity
(24-32 x 10-15 um; thallus pseudocyphellate) .......... Pyrenula sexlocularis
EASCOSPOLES SSS CP UAC Ce 8 rr ene Perec cle eeenah oO sare 2 sagt er tensa A cat tenant! Scat 24
Ascomata mostly aggregate with fused walls but with separate ostioles
(pseudostromatic) (ascospores 15-20 x 6-8 um) ........... Pyrenula anomala
AScommata-TNOStLy SIE aca 4 ee 8's cpt beh Stes eto ee Reo Bt oe Ltt lo 25
Thallus ecorticate (whitish; ascospores 17-25 x 7-11 um) .. Pyrenula microcarpa
Dallas CQrtica tes sat te aB dM nal PRs seal PS ent Peden at staal Pee, Re ella 2, «dig 26
Terminal lumina directly against the exospore ............ 0.0... e eee eee eee 27
Terminal lumina separated from exospore by thickened endospore wall........ 28
Hamathecium inspersed with oil droplets;
ascospores 16-24 x 8-13 UM ..... eee eee eee ee Pyrenula fetivica
Hamathecium clear; ascospores 17-22 x 9-12 um ............ Pyrenula nitidula
Hamathecium inspersed with oil droplets ........ 2... cece eee eee eee eee 29
Famiatheciiincleate... AA ht eter ell rk, all ork will ark tal ark ealt tak gat a cial at 32
ASconiata-S0. 741M, Wy CISA. Lae ee Rabie ake Sader Baden alee Dall beta arte 30
Aseomatas@.7minPidiay, 5 2 i Fes 4 Rt on Sy oy ee Ee ee elas Rene Pains 31
Ascospores 13-15% 4=6, UM +2. lexi ese ns eae wis wings sees x Pyrenula laetior
ASCOSPOTES G22 Be Voi days cg hae neg f fel een fee es Pyrenula subglabrata
JAscospores 15220 KA 9575 (ID, hi ali aan vial aa posh ahs whe Pyrenula mamillana
PASCOSPOLES S225. OF) WY. hance Bore Paks a Pol oe Pokese Pyrenula massariospora
AsScospores anostlyee2S wind LONG: 10a cw Wea oe weer eae yan nee age ae. 33
Ascospores. mostly <25.utw longs 25 .e ces edee a ebae 5 eegt yeh a vlan Bs eyed ving 34
Thallus pseudocyphellate; ascomata immersed ............. Pyrenula quassiicola
Thallus lacking pseudocyphellae; ascomata emergent ...... Pyrenula mastophora
Ascospores mostly 21-25 um long (ascomata <0.5 mm diam.) ... Pyrenula nitidella
ASCOSDONES 21 DM IONG or a heuste at Melo hehe one Aeon Macatee ane Ate cake ons 35
AS COSMOFES OST AUS: Ma LST og, Shin anche nstca sh vondsalt- nonstate wench pendteade wom 36
ASCOSPOTES DIOS SU SAT LOIN Se on «ia ts > fb He or a bes rca He Pes a MG bra ty begs bk: Sng he! 37
ASCOSpores ASG WIE sh souls oud fn eur swe smaols abou sn De Pyrenula aspistea
Ascospores 6=8 ttt Wide «4... ne nce s cone ee ctewe water asinns Pyrenula brunnea
Ascomata 0.4-0.7 mm in diam. ............... 2c eee eee ee Pyrenula aggregata
AS COMM ALA cee HD PAMELA Yep act: Or ne A non tbaugi tte Nart eget aedingise OE eeeeae eee os 38
Lumina round, ascospores 18-22 x 6-9 UM ......... ee eee eee Pyrenula scutata
Lumina angular, ascospores 19-22 x 7-8um ..............0006. Pyrenula balia
Ascomata mostly aggregated with shared ostiole (astrothelioid)
(ascospores 100-132 x 32-42 um) .............. Anthracothecium interlatens
AS COMMALAAIIOSUY CCUG O Png ucccehacd net Pn ounase dS nonca dst blest Mt tafe peop Se yee 40
968 ... Joshi, Upreti & Hur
40. Ascospones <200 tin lone 92.8, ot 45, oI AS aah tae ta as hee 4]
AO SASCOSPOTes STOO Mie ONS rt oe nsssasetots eae e nee Mimo kate easter Banjos 8 Me 47
41. Thallus and ascomata with orange-yellow anthraquinone
fascospores.12-25.« 8-131) 6 ello la atin as Pyrenula ochraceoflava
41. Thallus and ascomata lacking anthraquinone ................ 0... c ee eee eae 42
42. Ascospores 12-23 x 8-12 UM... .. eee eee ee eee eee Pyrenula parvinuclea
AZ IASCOSDOLCS ZOU WINS hee a nat Pa pun PparaeeSe Paes SP Geaeee FP eee Pca AES ne «Ep ag 43
43. Mature ascospores filled with orange oil (38-50 x 15-20 um) ... .Pyrenula breutelii
43. Mature-ascospores lackins-orange oil... Poi le lp stgee fa sige bp sigea by digiat wratha S prs 44
44, Hamathecium inspersed with oil droplets
(ascospores 30-45 um long) ............. 0. eee eee eee Pyrenula sublaevigata
HAS Tlatiatlveca unin, Cleats ¢ Csr 5 Aer. eet shane one SAM wna AM ana MA one eae OE are a 45
45. Ascospores 60-90 x 30-40 Um... eee eee ee ee eee Anthracothecium prasinum
Ay GCOS POLES So PMI! MOTB ws apa ne ern ha Ws m0 hts be Fos ya ts Preys He Fraga Ms bella th reese He eed abe | 46
46. Ascospores 33-45 x 11-16 UM... 2. eee eee ee eee Pyrenula thelemorpha
46. Ascospores 45-65 xX 15-25 UM... eee ee ee eee ee Pyrenula pyrenuloides
47. Thallus pseudocyphellate; ascospores 2 per ascus,
THEO 1S0 9 3045 ih th ee ee ela tars lh es Pyrenula duplicans
47. Thallus non-pseudocyphellate; ascospores 1-4 per ascus,
ZO S160 Se SOLAS Hite 14 et tern eae ee Anthracothecium macrosporum
Acknowledgments
This work was supported by a grant from the National Research Foundation of
Korea (NRF-2014K1A3A1A09063058) and the Korean National Research Resource
Center Program (NRF-2017M3A9B8069471). Santosh Joshi thanks Director, CSIR-
NBRI, Lucknow, India, for providing laboratory facilities. The authors are grateful
to Dr. André Aptroot (ABL Herbarium, The Netherlands) and Dr. Robert Lticking
(Botanischer Garten und Botanisches Museum, Berlin) for their valuable comments on
the manuscript.
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and Trypetheliaceae in the Malesian flora region. Blumea 54: 145-147.
https://doi.org/10.3767/000651909X475923
Aptroot A. 2012. A world key to the species of Anthracothecium and Pyrenula. Lichenologist
44: 5-53. https://doi.org/10.1017/S00242829 11000624
Aptroot A, Liicking R. 2016. A revisionary synopsis of Trypetheliaceae (Ascomycota:
Trypetheliales). Lichenologist 48: 763-982. https://doi.org/10.1017/S0024282916000487
Aptroot A, Sparrius LB. 2006. Addition to the lichen flora of Vietnam, with an annotated
checklist and bibliography. Bryologist 109: 358-371.
https://doi.org/10.1639/0007-2745(2006)109[358:ATTLFO]2.0.CO;2
Pyrenulaceae & Trypetheliaceae in Vietnam ... 969
Aptroot A, Licking R, Sipman HJM, Umafia L, Chaves JL. 2008. Pyrenocarpous lichens with
bitunicate asci. A first assessment of the lichen biodiversity inventory in Costa Rica. Bibliotheca
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Aptroot A, Ertz D, Etayo Salazar JA, Gueidan C, Mercado Diaz JA, Schumm F, Weerakoon
G. 2016. Forty-six new species of Trypetheliaceae from the tropics. Lichenologist 48:
609-638. https://doi.org/10.1017/S002428291600013X
Licking R, Nelsen MP, Aptroot A, Barillas De Klee R, Bawingan PA, Benatti MN, Binh
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LI, Grijalva A, Gueidan C, Hernandez M JE, Knight A, Lumbsch HT, Marcelli MP,
Mercado-Diaz JA, Moncada B, Morales EA, Naksuwankul K, Orozco T, Parnmen
S, Rivas Plata E, Salazar-Allen N, Spielmann AA, Ventura N. 2016. A phylogenetic
framework for reassessing generic concepts and species delimitation in the lichenized
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https://doi.org/10.1017/S0024282916000505
Orange A, James PW, White FJ. 2010. Microchemical methods for the identification of
lichens. British Lichen Society.
Upreti DK. 1991. Lichen genus Pyrenula from India: the species with spores of Pyrenula
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241-247. https://doi.org/10.1080/01811797.1991.10824926
Upreti DK. 1998. A key to the genus Pyrenula from India, with nomenclatural notes. Nova
Hedwigia 66: 557-576.
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 971-975
https://doi.org/10.5248/132.971
Acumispora delicata sp. nov.
from the Brazilian Atlantic Forest
PHELIPE M.O. CosTA', MARCELA A. BARBOSA’, WANDERSON L. TAVARES’,
DAYNET SOSA?*, SIMON PEREZ-MARTINEZ?+, RAFAEL F. CASTANEDA-RUIZ?
& ELAINE MALOSSO”
‘Centro de Biociéncias, Departamento de Micologia
Universidade Federal de Pernambuco, Avenida da Engenharia,
s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil
? Programa de Pés-Graduag¢do em Biologia de Fungos,
Universidade Federal de Pernambuco, Avenida da Engenharia,
s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil
> Escuela Superior Politécnica del Litoral, ESPOL, (CIBE), Campus Gustavo Galindo,
Km. 30.5 Via Perimetral, P.O. Box 09-01-5863, Guayaquil, Ecuador
‘Universidad Estatal de Milagro (UNEMI), Facultad de Ingenieria, Cdla. Universitaria,
Km. 1.5 Via Milagro-Km26, Milagro 091706, Guayas, Ecuador
° Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT)
Alejandro de Humboldt, OSDE, Grupo Agricola,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, C.P. 17200, Cuba
* CORRESPONDENCE TO: elaine.malosso@ufpe.br
ABSTRACT—A new species, Acumispora delicata, collected on decaying leaves of an
unidentified plant, is described and illustrated. The new species is distinguished by
narrow ovoid to broad obclavate conidia, which are acuminate toward the rounded or
obtuse apex. A key and illustrations of Acumispora species are provided.
Key worps—asexual fungi, neotropic, taxonomy, systematics
Introduction
The genus Acumispora was described by Matsushima (1980) and typified by
Acumispora uniseptata Matsush. It is characterized by conidiophores that are
macronematous or micronematous, unbranched or irregular branched, mostly
prostrate, sometimes erect, septate, brown or pale brown, sometimes reduced
872 ... Costa & al.
to conidiogenous cells that are monoblastic, determinate or polyblastic, after
several sympodial extensions, integrated or discrete, denticulate. The conidia
are solitary, acrogenous or acropleurogenous, navicular, narrow ovoid,
obclavate, acuminate or rostrate toward the apex, very pale brown to brown,
with a minute basal frill after the rhexolytic conidial secession.
During a mycological survey of microfungi associated with leaf litter in a
Brazilian Atlantic forest, a conspicuous fungus was collected. It is described as
a new Acumispora species.
Materials & methods
Individual collections were placed in plastic bags, taken to the laboratory, and
treated according to Castafieda-Ruiz et al. (2016). Mounts were prepared in polyvinyl
alcohol-glycerol (8 g PVA in 100 ml water, plus 5 ml glycerol) and lactofuchsin
(0.1 g acid fuchsin, 100 ml 85% lactic acid) following Carmichael (1955) or lactic acid
(90%). Measurements were made at a magnification of x1000 under a Nikon Eclipse
Ni-U microscope with bright field optics, and photomicrographs were taken using DIC
optics with a Nikon DS-Fi2 camera. The holotype was deposited in the Herbarium of
Universidade Federal de Pernambuco, Recife, Brazil (URM).
Taxonomy
Acumispora delicata P.M.O. Costa, Malosso & R.E. Castafieda, sp.nov. —_- Fics 1, 2B
INDEX FUNGORUM IF 554059
Differs from Acumispora biseptata by its smaller, 3-septate conidia.
Type: Brazil, Pernambuco, Igarassu, RPPN Reftigio Ecolégico Charles Darwin, 7°48’S
34°57’W, on decaying leaves of an unidentified plant, 12.V.2017, coll. P.M.O. Costa
(Holotype, URM 90134).
ErymMo_oey: Latin, delicata, means delicate, elegant, graceful.
Co.Lonigs on the natural substrate, epiphyllous, slightly funiculose, scattered
granulose, yellowish-brown. Mycelium mostly superficial, composed of septate,
unbranched, pale yellowish-brown, smooth-walled hyphae, 1-2 um diam.
CONIDIOPHORES micronematous, prostrate, mostly reduced to conidiogenous
cells. CONIDIOGENOUS CELLS monoblastic, discrete, determinate, lageniform,
abruptly tapered, cylindrical at the conidiogenous loci, pale yellowish-brown to
pale brown, 4-8 x 3-3.5 um. Conidial secession rhexolytic. Conip1a solitary,
acrogenous, narrow ovoid to broad obclavate, acuminate toward the rounded
or obtuse apex, straight, 3-septate, smooth, pale yellowish-brown or pale
brown, 15-20 x 3-4 um, dry.
Notes: Four species have been accepted previously in Acumispora:
A. biseptata Matsush., A. phragmospora Matsush., A. wuniseptata, and
A. verruculosa Heredia et al. (Heredia et al. 2007, Matsushima 1980). Acumispora
Acumispora delicata sp. nov. (Brazil) ... 873
Fic. 1. Acumispora delicata (ex holotype, URM 90134). A. Conidia. B. Conidiogenous cells.
delicata resembles A. biseptata, which differs by its 2-septate, constricted at the
septa, longer (16-25 um) conidia, with a triangular, elongated, rostrate apical
cell. Conidia of the five species are illustrated in Fic. 2.
Key to Acumispora species
Tee ON IEA ATTAO O UNS CMe fo Mee oo MOe oO ins Mee ine yO ane we ate Aen Ieee ede eee 2
1. Conidia verruculose (obclavate to slightly ovate, 2-septate,
constricted at the septa, 25.4-33.5 x 4.4-5.5 um) ............0.. A. verruculosa
2. Conidia 1-septate (inequilateral, more or less navicular to slightly obturbinate,
with linear-triangular, rostrate apical cell, 23-28 x 5.5-6.5 um) ... A. uniseptata
2 C otiidia with more than One Sept: os. 8h os asta ib eats a gee ares 3
3. Conidia 2-septate (narrow ovoid with linear-triangular elongated,
rostrate apical cell, 16-25 x 4-5 um) ...... eee eee eee ee eee A. biseptata
3: Conidiasyith-more than 2-septates st hs ot tye ty dette $e de Bike Lhe +
4. Conidia 3-septate, narrow ovoid to broad obclavate,
acuminate toward the apex, 15-20 x 3-4um ................0.06. A. delicata
4, Conidia 3-6-septate, cylindrical-fusiform or long navicular,
apexacuiminates 20235 MAH SIM: 2h caw sh pe een zee eed A. phragmospora
874 ... Costa & al.
A
£ £
= =
Oo °
D
C =
=
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= -
=
O
Fic. 2. Acumispora spp., conidia: A. A. biseptata (Matsushima 1980); B. A. delicata (ex holotype,
URM 90134); C. A. phragmospora (Matsushima 1980); D. A. uniseptata (Matsushima 1980);
E. A. verruculosa (Heredia et al. 2007).
Acumispora delicata sp. nov. (Brazil) ... 875
Acknowledgments
The authors express their sincere gratitude to Dr. Xiu-Guo Zhang and Dr. De-Wei Li
for their critical review of the manuscript. The authors are grateful to the Coordenacao
de Aperfeigoamento de Pessoal de Nivel Superior (CAPES) for financial support through
Project 88881.062172/2014-01 and the Programa Ciéncia sem Fronteiras. RFCR is
grateful to the Cuban Ministry of Agriculture and Programa de Salud Animal y Vegetal,
Project P131LH003033 for facilities. DS is grateful to the Escuela Superior Politécnica
del Litoral, ESPOL, (CIBE) for financial support. We acknowledge the facilities provided
by Dr. P.M. Kirk and Drs. V. Robert and K. Bensch, through the Index Fungorum and
MycoBank websites. Dr. Lorelei Norvell’s editorial review and Dr. Shaun Pennycook’s
nomenclature review are greatly appreciated.
Literature cited
Carmichael JW. 1955. Lacto-fuchsin: a new medium for mounting fungi. Mycologia 47: 611.
Castafieda-Ruiz RF, Heredia G, Gusmao LFP, Li DW. 2016. Fungal diversity of Central and
South America. 197-217, in: DW Li (ed.). Biology of Microfungi, Springer International
Publishing. https://doi.org/10.1007/978-3-319-29137-6_9
Heredia G, Castafieda-Ruiz RF, Arias RM, Saikawa M, Stadler M. 2007. Anamorphic fungi from
submerged plant material: Acumispora verruculosa sp. nov., Pleurophragmium aquaticum sp.
nov and Pleurophragmium miniumbonatum comb. nov. Mycotaxon. 101: 89-97.
Matsushima T. 1980. Saprophytic microfungi from Taiwan. Part. 1. Hyphomycetes. Matsushima
Mycological Memoirs No. 1. 82 p.
MY COTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October-December 2017— Volume 132, pp. 977-983
https://doi.org/10.5248/132.977
Anisogenispora insignissima gen. & sp. nov.
from the Brazilian semi-arid region
SHEILA MIRANDA LEAO-FERREIRA’,
Luis FERNANDO PASCHOLATI GUSMAO! & RAFAEL F. CASTANEDA-RUIZ?
' Universidade Estadual de Feira de Santana
Avenida Transnordestina, s/n, Novo Horizonte, 44036-900, Feira de Santana, Brazil
? Instituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT) Alejandro de Humboldt,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
* CORRESPONDENCE TO: Igusmao@uefs. br
ABSTRACT—A new genus and species Anisogenispora insignissima is described and illustrated.
The fungus, found on decaying fruit of an unidentified dicotyledonous plant, is distinguished
by blastic production of globose to broadly pyriform, 1-2-septate, bicolored conidia and
a Thielaviopsis-like synanamorph with lenticular, doliiform, cylindrical, or vermiform,
unicellular or multiseptate, black conidia produced by thallic-arthric disarticulation. All
conidiogenous events originate on the same conidiophores.
KEY worDs—asexual fungi, hyphomycetes, taxonomy, tropics
Introduction
The Brazilian semiarid region, located in the northeast of the country,
covers approximately 900,000 km? and exhibits a high biodiversity (Giulietti
et al. 2006). Its different vegetation has favored highly diverse microfungi and
the discovery of several new species (e.g., Almeida et al. 2014, Barbosa et al.
2013, Fiuza et al. 2014, Leao-Ferreira et al. 2015, Monteiro et al. 2016, Silva &
Gusmao 2013, Silva et al. 2014). During a mycological survey of conidial fungi
from the semi-arid region in Serra do Ramalho, Bahia State, an interesting
fungus was collected that showed remarkable differences from all previously
described hyphomycetes (Seifert et al. 2011); it is described here as a new genus
and species.
978 ... Leao-Ferreira, Gusmao & Castafieda-Ruiz
Materials & method
Samples of decaying plant materials collected in Serra do Ramalho, Bahia State
were placed in plastic bags for transport to the laboratory where they were treated
according to Castafieda-Ruiz et al. (2016) and placed in humid chambers. Several
attempts to obtain these species in pure culture were unsuccessful after transferring
conidia with a flamed needle to malt extract agar [2% (w/v) malt extract and corn meal
agar, mixed 1:1 with carrot extract] incubated at 25 °C. Mounts were prepared in PVL
(polyvinyl alcohol, lactic acid) and measurements were made at 1000x magnification.
Microphotographs were obtained with an Olympus BX51 microscope equipped with
bright field and Nomarski interference optics. The type specimen is deposited in the
Herbarium of Universidade Estadual de Feira de Santana, Bahia, Brazil (HUEFS).
Taxonomy
Anisogenispora S.M. Leao, Gusmao & R.F. Castaneda, gen. nov.
INDEX FUNGORUM IF 552545
Differs from Repetophragma by its monoblastic and polyblastic conidiogenous cells with
percurrent and sympodial extensions and by its chlamydospore-like synanamorph with
thallic-arthric disarticulation originating near the conidiophore apex.
TYPE SPECIES: Anisogenispora insignissima S.M. Leao et al.
ETYMOLOGY: aniso- (Greek) meaning unequal, uneven, or dissimilar + -geni- (Latin)
meaning born or produced and referring to the blastic and thallic modes of conidial
ontogeny + -spora (Latin) referring to the conidia.
CoLonies on the natural substrate effuse, brown. CONIDIOPHORES
macronematous, mononematous. CONIDIOGENOUS CELLS monoblastic
and polyblastic, integrated, indeterminate, with enteroblastic percurrent
extension, sometimes polyblastic with sympodial extension; conidial secession
schizolytic. Conrp1A solitary, brown, euseptate, smooth, dry. SYNANAMORPH
Thielaviopsis-like, terminal, dark brown, euseptate, originating near the apex of
the conidiophores, forming conidia by thallic-arthric disarticulation; conidial
secession schizolytic; conidia unicellular or multicellular, dark brown or black,
smooth.
Anisogenispora insignissima S.M. Leao, Gusmao & R.F. Castafieda, sp. nov. Fics 1-3
INDEX FUNGORUM IF 552546
Differs from Repetophragma biseptatum and R. quadriloculare by its conidiogenous cells
with percurrent and sympodial extension and by its Thielaviopsis-like synanamorph
originating from the conidiophores and conidia.
Type: Brazil, Bahia State, Coribe, Serra do Ramalho, 43°47’W 13°32’S, alt. 450-600 m,
on fallen decaying fruit of an unidentified dicotyledonous plant, 16.11.2008, coll. S.M.
Leao-Ferreira (Holotype, HUEFS 210449).
ETYMOLOGy: insignissima (Latin) meaning very remarkable, referring to pleomorphic
conidia produced on the same conidiophores.
Anisogenispora insignissima gen. & sp. nov. (Brazil) ... 979
B
Fic. 1. Anisogenispora insignissima (holotype, HUEFS 210449). A-C. Conidia. D, E. Conidiogenous
cells and conidia. EF Conidiophores, conidiogenous cells, and conidia. Scale bars = 10 um.
980 ... Ledo-Ferreira, Gusmao & Castafieda-Ruiz
CoLonigs on the natural substrate effuse, hairy-granulose, dark brown to
black. Mycelium superficial and immersed, composed of septate, branched,
smooth, 2-4 um diam, brown to pale brown hyphae. CoNnIDIOPHORES
macronematous, mononematous, erect, flexuous, straight or geniculate,
unbranched, rarely branched, 2-7-septate, brown, or alternately and irregularly
versicolored brown, pale brown or dark brown, 90-300 x 7.5-10 um, with
4-12 enteroblastic percurrent extensions, smooth. CONIDIOGENOUS CELLS
monoblastic, sometimes polyblastic, cylindrical, integrated, indeterminate,
terminal becoming intercalary with frequent and numerous enteroblastic
percurrent extensions and sometimes with holoblastic, sympodial extension
after a percurrent extension, 8-20 x 3 um, smooth, pale brown to brown.
CONIDIOGENOUS LOCI flat. Conip1a solitary, acrogenous or acropleurogenous,
broad pyriform, subglobose, somewhat turbinate to obovoid, (1-)2-septate,
bicolored, basal and apical cells pale brown, middle cell dark brown or black,
smooth, 17.5-30 x 12.5-20 um. SyNaANAMORPH Thielaviopsis-like, single,
doliiform, cylindrical, vermiform, or long fusiform to navicular, mostly
terminal, 2-16-euseptate, slightly constricted at the septa, 50-200 x 15-28 um,
very dark brown or black, originated near or at the apex of the conidiophores
and conidia, forming thallic-arthric conidia by disarticulation at the septa;
conidial secession schizolytic; conidia lenticular, doliiform, cylindrical,
0-12-septate, dark brown or black, 8-120 x 13-28 um, smooth, dry.
Note: Anisogenispora insignissima resembles Repetophragma biseptatum
(M.B. Ellis) Subram. and R. quadriloculare (Matsush.) R.F. Castafieda et al. in
conidial shape and the enteroblastic percurrent extension of the conidiogenous
cells, but R. biseptatum and R. quadriloculare do not have sympodial extension
of the conidiogenous cells (Castafeda-Ruiz et al. 2011), nor do they produce
a Thielaviopsis-like synanamorph with thallic-arthric conidial ontogeny such
as is present on the conidiophores in A. insignissima. Hyphae in some species
of Chalara (Corda) Rabenh. develop Thielaviopsis-like synanamorphs (similar
to the A. insignissima synanamorph), but Chalara exhibits an enterogenous
conidial ontogeny, considered as phialidic (Nag Raj & Kendrick 1976). The
Thielaviopsis-like synanamorph produced in Arthrotaeniolella L.B. Conc. et al.
Fic. 2. Anisogenispora insignissima (holotype, HUEFS 210449). A. Young growing conidium
from Thielaviopsis-like synanamorph after its germination. B. Conidiogenous cells, conidia, and
Thielaviopsis-like synanamorph. C,D. Thallic-arthric conidia of Thielaviopsis-like synanamorph.
E, FE. Conidiogenous cells, conidia, and thallic-arthric conidia of Thielaviopsis-like synanamorph.
Scale bars = 10 um.
Anisogenispora insignissima gen. & sp. nov. (Brazil) ... 981
982 ... Leao-Ferreira, Gusmao & Castafieda-Ruiz
Fic. 3. Anisogenispora insignissima (holotype, HUEFS 210449). Conidiophores, conidiogenous
cells, and conidia; and thallic-arthric conidia of Thielaviopsis-like anamorph.
is also somewhat similar to that of A. insignissima, but Arthrotaeniolella species
are freshwater fungi with thallic-arthric disarticulation of the conidiophore
branches (Monteiro et al. 2017).
Anisogenispora insignissima gen. & sp. nov. (Brazil) ... 983
Acknowledgments
We are indebted to Dr. Xiu Guo Zhang and Dr. De-Wei Li for their critical reviews.
The authors thank to the National Council for Scientific and Technological Development
(CNPq; Proc. 451991/2016-8). The first author is grateful to Coordination for the
Improvement of Higher Education Personnel (CAPES) through project N° 071/2012.
RFCR is grateful to OSDE, Grupo Agricola from the Cuban Ministry of Agriculture and
Programa de Salud Animal y Vegetal, project P131LH003033. Dr. Lorelei L. Norvell’s
editorial review and Dr. Shaun Pennycook’s nomenclature review are greatly appreciated.
Literature cited
Almeida DAC, Miller AN, Gusmao LFP. 2014. New species and combinations of conidial
fungi from the semi-arid Caatinga biome of Brazil. Nova Hedwigia 98: 431-447.
https://doi.org/10.1127/0029-5035/2013/0162
Barbosa FR, Raja HA, Shearer CA, Gusmao LFP. 2013. Some freshwater fungi from the Brazilian
semi-arid region, including two new species of hyphomycetes. Cryptogamie, Mycologie 34:
243-258. https://doi.org/10.7872/crym.v34.iss2.2013.243
Castaneda-Ruiz RF, Heredia G, Arias RM, McKenzie EHC, Hyde KD, Stadler M, Saikawa M, Gené
J, Guarro J, Iturriaga T, Minter DW, Crous PW. 2011. A new species and re-disposed taxa in
Repetophragma. Mycosphere. 2: 273-289.
Castafieda-Ruiz RF, Heredia G, Gusmao LFP, Li DW. 2016. Fungal diversity of Central and South
America. 197-217, in: DW Li (ed.). Biology of Microfungi. Springer International Publishing.
https://doi.org/10.1007/978-3-319-29137-6_9
Fiuza PO, Gusmao LFP, Cruz ACR, Castaneda Ruiz RF. 2014. Conidial fungi from the semiarid
Caatinga biome of Brazil: a new species of Pseudoacrodictys. Mycotaxon 127: 33-37.
https://doi.org/10.5248/127.33
Giulietti AM, Harley RM, Queiroz LP, Rapini A. 2006. To set the scene. In: Giulietti AM, Queiroz
LP, Rapini A. (Eds). Towards greater knowledge of the Brazilian semi-arid biodiversity.
Ministério da Ciéncia e Tecnologia. Brasilia.
Ledo-Ferreira SM, Gusmao LFP, Almeida DAC, Castafeda- Ruiz RE 2015. Digicateno-
sporium polyramosum, a new hyphomycete from Brazil. Mycotaxon 130: 479-482.
https://doi.org/10.5248/130.479
Monteiro JS, Gusmao LFP, Castafeda-Ruiz RF. 2016. Pleurothecium bicoloratum & Sporidesmiopsis
pluriseptata spp. nov. from Brazil. Mycotaxon 131: 145-152. https://doi.org/10.5248/131.145
Monteiro JS, Gusmao LFP, Conceicéo LB, Castafeda-Ruiz RE 2017. Arthrotaeniolella aquatica
gen. & sp. nov. and Pseudospiropes piatanensis sp. nov. from Brazil. Mycotaxon 132: 373-379.
https://doi.org/10.5248/132.373
Nag Raj TR, Kendrick WB. 1976. A monograph of Chalara and allied genera. Wilfrid Laurier
University Press, Waterloo, Ontario, Canada. 200 p.
Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS
Biodiversity Series 9. 997 p.
Silva SS, Gusmao LFP. 2013. Conidial fungi from the semi-arid Caatinga Biome of Brazil. A new
species of Dictyochaeta. Mycosphere 4(4): 701-705. https://doi.org/10.5943/mycosphere/4/4/6
Silva SS, Cruz ACR, Gusmao LFP, Castafeda-Ruiz RF. 2014. Diplococcium variegatum, a new
conidial fungus from the semi-arid Caatinga biome of Brazil. Mycotaxon 127: 59-62.
https://doi.org/10.5248/127.59
MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. © 2017
October-December 2017— Volume 132, pp. 985-986
https://doi.org/10.5248/132.985
Regional annotated mycobiotas new to the Mycotaxon website
ABSTRACT—Mycotaxon is pleased to announce two new species distribution lists to our
‘web-list’ page covering diversity of wood-inhabiting aphyllophoraceous basidiomycetes
on the island of Cyprus (by Loizides) and saprobic asexual microfungi from the tropical
cloud forest of Veracruz, México (by Arias, Heredia & Castafieda-Ruiz). This brings
to 127 the number of free access mycobiotas now available on the Mycotaxon website:
http://www.mycotaxon.com/resources/weblists.html
EUROPE
Cyprus
MicHaEt Loizipes. Diversity of wood-inhabiting aphyllophoraceous
basidiomycetes. 27 p.
ABSTRACT—The diversity of wood-inhabiting aphyllophoraceous basidiomycetes
on the island of Cyprus is explored in this paper, following a ten-year inventory
between 2007 and 2016. A total of one-hundred-and-eight taxa are reported,
fifty-eight of which constitute new records for the country. Twenty-two species
occurring on the narrow-endemic golden oak of Cyprus (Quercus alnifolia) are
documented for the first time on this host. Collections of Laetiporus sulphureus
sensu lato from Ceratonia and Eucalyptus are phylogenetically analysed and
revealed to belong to a distinct clade likely representing an undescribed species.
Among the newly reported taxa, of particular interest are Amaurodon viridis,
Asterostroma ochroleucum, Byssomerulius hirtellus, Crustoderma dryinum,
Dendrocorticium polygonioides, Postia inocybe, P. simani, Steccherinum ciliolatum,
and S. oreophilum, all of which are considered rare and are seldom reported
in literature. The aggressive conifer pathogens Heterobasidion annosum and
Porodaedalea pini are also rare and do not appear to have a significant impact
on the island’s pine-dominated forests. Fuscoporia torulosa, on the other hand,
is commonly encountered on sclerophyllous vegetation and should be closely
monitored. Previous aphyllophoraceous records from the island are critically
discussed and re-evaluated, with old and new data compiled in the form of an
annotated checklist, to include notes on the substrate, fruiting season, altitude,
and estimated abundance.
986 ... New regional mycobiotas online
NORTH AMERICA
Mexico
Rosa Maria ARIAS, GABRIELA HEREDIA & RAFAEL F. CASTANEDA-
Ruiz. Checklist of saprobic asexual microfungi from the tropical
montane cloud forest of Veracruz, México. 48 p.
Axsstract— A checklist comprising 355 species and 190 genera of asexual
microfungi associated with plant debris collected in different localities of
the tropical montane cloud forest of Veracruz, Mexico, is presented based on
literature and fresh specimens. Annotations (lists of studied material and
locations) are accompanied by illustrations of 31 species. A total 101 taxa are
recorded for the Mexican mycobiota for the first time.
MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. © 2017
October-December 2017— Volume 132, pp. 987-1001
https://doi.org/10.5248/132.987
BOOK REVIEWS AND NOTICES.
LORELEI L. NORVELL & ELSE VELLINGA, EDITORS
Pacific Northwest Mycology Service, Portland OR 97229-1309 USA
ABSTRACT—Books reviewed include: BasiplomyceTES—Agaricus of North America
(Kerrigan 2016), Hebeloma (Fr.) P. Kumm. (Beker et al. 2016); LrcHens—Lichens of
Mexico (Herrera-Campos et al. 2016).
BASIDIOMYCETES
Agaricus of North America. By Richard W. Kerrigan. 2016. Memoirs OF THE
NEw YorRK BOTANICAL GARDEN, VOL. 114. NYBG Press, Bronx NY 10458-5126, USA.
592 p. ~385 color plates & figures, hardback. ISSN 0077-8931; ISBN 978-0-89327-536-5.
Price (excl. postage): $127.99. www.nybgpress.org
c das
; i praeclaresquamosus is chewy
But the name here turns out to be hooey,
See ic - : And those who like poison
AG AREC US . Will now find their joys in
of NORTH AMERICA, | Agaricus buckmacadooi.
. If you take an Agaricus census,
RICHARD W. KERRIGAN
7
The naming will batter your senses,
Some mushrooms that pose as
‘praeclaresquamosus’
Should really be deardorffensis.
The above limerick, dashed off today by British
Columbian Ian Gibson (our Pacific Northwest
Key Council database expert), captures
the west coast’s response to Rick Kerrigan's
new masterpiece. A trained agaricologist,
* Book reviews or books for consideration for coverage in this column should be sent to the
Editor-in-Chief <editor@mycotaxon.com> 6720 NW Skyline, Portland OR 97229 USA.
988 ... MycoTAxon 132(4)
I nonetheless still embarrass myself hemming and hawing over a pristine
Agaricus presented for field identification. Confidently identifying Inocybe,
Phaeocollybia, Chrysomphalina, Galerina, and many small or ‘valueless’
mushrooms, I am flummoxed by the genus that gave agaricology its name.
Somewhat defeated by identifications that are misapplications of European
concepts, most North American field agaricologists slavishly follow Arora
(1986) who—recognizing an Agaricus identification crisis when he saw one—
cheerfully placed ‘group’ after species names to give the uncertain identifier a
bit of wiggle room. But I digress.
AGARICUS OF NorTH AMERICA is a sturdy hefty tome with stiff hard covers
designed to stand up to frequent use. Owners are nonetheless advised to
prepare the volume by splaying it firmly flat in sections to prime it for heavy
use on lab benches.
Presentation is thorough and comprehensive. The table of contents and
acknowledgments precede introductory chapters covering Agaricus in the
(1) world and (2) commerce; history & nomenclature; literature; “To study
and identify Agaricus” (a materials & methods section to read and re-read);
formats & conventions; edibility caveats; phylogenetic analysis; sequences
representing Agaricus sections (+ subsections & unnamed major lineages);
and typifications of 10 previously published taxa.
These introductory chapters provoke. In “Why this book? Why now?”
Kerrigan does not share his 45 years of Agaricus field experience in a field guide
because “I believe that this presentation uniquely bridges a gap between the
typical specialist literature and the typical field guide in a way that will interest
many readers.” He further notes—
“What I have attempted to produce is a MONOGRAPHIC TECHNICAL
RESOURCE that is authoritative, while remaining generally accessible,
and is comprehensive to the extent that our existing knowledge allows.
... Agaricus is a difficult, subtle, phenotypically plastic genus with many,
many rare and seemingly cryptic species, and North America is a big
and diverse continent. ... The present volume ... is just a start, a down
payment on what we as stewards of the commons deserve: an exhaustively
comprehensive, absolutely authoritative summary of Agaricus in North
America.... The process is far from completed.”
The author ends his opening chapter with the challenge—“See if you can
discover what I’ve missed.”
In “Agaricus erected, disused, emended, and conserved” Rick observes—
“As [Linnaeus] employed the name, Agaricus corresponded to the gilled
mushrooms. Consequently, for more than a century, most new mushrooms
were named as an Agaricus, creating a huge list of names for a diverse
Book Reviews ... 989
range of species (now mostly reassigned to many segregated genera). This
copious nomenclatural residue is just one more cumbersome obstacle,
virtually unique to Agaricus, impeding the simple use of correct names,
thus affecting comprehension and communication.”
The 33-page chapter on how to approach the genus clearly explains why so
many of the collections studied are Kerrigan’s own and how to avoid making
collections that are destined to reside only in herbaria—uncharacterized,
unattended, and decidedly unloved. The helpful five pages devoted to the
objective of Agaricus study [“I have applied the names that in my opinion
are MOST PROBABLY correct’] precede discussions on characters (macro-/
microscopic, eco-/geological, biochemical, molecular, cultural), all summarized
in a brief paragraph on p. 49.
Next follow the all-important keys and taxonomic treatments. ‘The titles
explain what the keys are designed to do and not do: [i] “Dichotomous key for
the placement of agaricoid and secotioid specimens of Agaricus subg. Agaricus
in North America to sections or groups” and “[ii] Quasi-synoptic key for the
placement of specimens of A. subg. Agaricus in North America to sections or
groups.” The author shares his frustration in crafting keys: “..natural groups
in Agaricus...are somewhat heterogeneous morphologically and sometimes
biochemically, and frustrate attempts to produce elegant dichotomous keys,
especially when emphasizing field characters.” Hence the synoptic key serves as
“the preferred framework for a holistic assessment of attributes and affinities.
Once a section or group is selected, one or more subsidiary dichotomous
keys for the individual sections follow as each section is introduced....” One
complaint may be raised at using only the rDNA ITS region to support new
phylogenetic sections, and readers are warned to expect several taxonomic
shifts after future multigene sequence analyses are employed.
The formal taxonomic section covers 190 taxa (some represented only
by RWK collection numbers) and proposes 34 new names. Descriptions are
ordered based on formal section (Bivelares, Chitonioides, Xanthodermatei,
Sanguinolenti, Nigrobrunnescentes, Agaricus, Spissicaules, Subrutilescentes,
Rarolentes, Minores, Arvenses) (further divided into subsections) or the informal
‘floridianus, ‘longuloid/gryophragmioid, and ‘martineziensis’ lineages.
A typical section (e.g., A. sect. Bivelares) begins with a standard nomenclator
(but containing only selected synonyms) followed by AFFINITIES, CHARACTERS,
the inevitably valuable REMARKS, and KEY to species. This may or may not
precede SUBSECTIONS (with their own CHARACTERS, REMARKS, and KEY). Each
section/subsection includes an illustration, a graph comparing spore sizes, and
a phylotree.
990 ... MycoTaxon 132(4)
One to two illustrations accompany each 1-4-page species treatment: most
are photos—some in situ, most ‘morgue’ shots of fresh material (so useful
to taxonomists but too often snubbed by those hoping to see mushrooms in
natural habitats), and several of herbarium exsiccati—with a few drawings or
paintings. For reasons obvious to those who know how few Agaricus features
are diagnostic under high power, there are precious few micrographs.
Species sections include nomenclator, AFFINITIES; technical DESCRIPTION;
CHEMISTRY; HABIT, HABITAT, DISTRIBUTION; an always insightful piscussi1on;
and EpIBILiTy. [Edibility for Agaricus bisporus is noted as: “This mushroom
is a prized edible, the most commonly cultivated mushroom in the world,
the basis of a multi-billion-dollar industry. Cultivation is not difficult—if you
know how.’] The amount of new information (based on a lifetime of personal
observation) presented here is awe-inspiring.
And yet, there are big swaths of North America that are only superficially
studied (the southern and southeastern states), while Mexico merits only a
brief mention in the literature section. Most material in the book was collected
in California, Pennsylvania, Colorado, and New Mexico. Focused on one
continent as it is, the current volume presents keys and descriptions to cover
only species confirmed by the author as occurring there.
Since the 1970s, North American agaricologists have confronted far
too many misapplied names and a somewhat tumultuous “nomenclatural
inconsistency” in the continent’s monographs and field guides. As a test,
I consulted the 55 names listed in the PNW Key Council provisional key to
Pacific Northwest Agaricus species (Chariton 1997) to evaluate names accepted
by Kerrigan against those still used by most North American field taxonomists.
First eliminated were 18 provisional names from an unpublished master’s thesis
(Isaacs 1963); based on Oregon and Washington specimens, these were never
formally proposed and thus (with one exception) missing from Kerrigan's
index. After removing synonyms and names with no modern concept and
adding nine newly named taxa, the PNW list tallied 39 Kerrigan-accepted
Agaricus names. That list will grow once the PNW Key Council further mines
the volume to identify which additional species inhabit British Columbia,
Idaho, western Montana & Alberta, Washington, and Oregon.
The verses at the beginning encapsulate the problems faced by all
North American would-be Agaricus identifiers. In 1972, I identified one
forest mushroom as Agaricus meleagris, which was ‘corrected’ in 1974 to
A. placomyces. In the early 80s, Oregon Mycological Society members
learned that the CORRECT west coast name was A. praeclaresquamosus
(Kerrigan 1986), only to find thereafter that it also would not serve:
Book Reviews ... 991
“The Agaricus meleagris-placomyces-praeclaresquamosus complex
continues to puzzle, although Rick Kerrigan and colleagues seem to have
successfully teased out the western North American cluster. After years
of my spelling A. praeclaresquamosus Freeman slowly for fellow field
mycologists, we now recognize the complex (in the Pacific Northwest at
least) as Agaricus moelleri Wasser” (Norvell 2016).
But wait! Now A. deardorffensis (which has sent people to the hospital in British
Columbia) and A. buckmacadooi (honoring Washington's own PNW Key
Councilor Buck McAdoo) replace the first four errant names (Kerrigan et al.
2005, Norvell 2017). Fortunately AGARICUS OF NORTH AMERICA elucidates
the misapplied and confused European and North American concepts; those
perplexed by this complex are well advised to read and commit to memory
pp. 141-143, even while being aware that that the deardorffensis ITS rDNA
(sequenced originally from that mushroom brought into the Vancouver
Emergency Room) differs in only ONE base pair from the Tibetan species,
A. tibetensis.
The dry humor permeating this scientific treatise on all things Agaricus
will reassure readers who want entertainment with their erudition. I was
particularly charmed by a note on the newly minted Agaricus julius Kerrigan
(p. 394): “MISAPPLIED NAMEs: It has been called “the Prince” (A. augustus)
in common usage in Colorado; perhaps now it will become the Emperor
formerly known as Prince.” Such wit is rarely encountered in nomenclators.
The volume concludes with biographies and Agaricus species of Kerrigan's
North American predecessors (Charles H. Peck, William A. Murrill,
Alexander H. Smith, Bill E Isaacs, Alice E. Freeman), literature cited, lists
of critically examined types and vouchers studied and/or sequenced and
accepted, nomenclatural citations for the covered taxa, and a 5-page index
inexplicably entitled ‘Index to scientific names now (or formerly) assigned to
Agaricaceae, with prior homonyms [perplexing in that a comprehensive list
from 1753 onwards might rival the current volume in length]. On the back of
the page containing Rick’s biography and photo is printed a 6-in/16-cm ruler,
an incitement to all readers to go outside and get busy.
Together with the European Agaricus tomes (Parra 2008, 2013) and the
ongoing work in Asia (e.g., Zhao et al. 2011), the genus is rapidly losing many
of its secrets. We note that the work by Parra (whom Kerrigan warmly credits
in his acknowledgments) is quite different in scope and execution than that
by Kerrigan. Not so surprising, as Europe has a much longer and more
thorough mycological tradition with many more active people in the field.
992 ... MYCOTAXON 132(4)
Kerrigan's solution to life’s Agaricus problems has been a long time coming
but does not disappoint. While taxonomic riddles remain (giving Rick more
to do in his off time), many Gordian knots have been sliced. Although we
still take refuge in Arora’s ‘groups, they are becoming smaller and more
approachable. Misery does love company, so perhaps most comforting of all
is that this North American Agaricus expert lets us know that he finds His
genus even more difficult than the rest of us do.
Arora D. 1986. Mushrooms demystified. (2"¢ ed.). Ten Speed Press, Berkeley.
959'p.
Chariton LR. 1997 (reformatted by Ian Gibson April 2003). Trial field key to
the species of Agaricus in the Pacific Northwest. Pacific Northwest Key
Council. < http://www.svims.ca/council/Agari2.htm>
Isaacs BF. 1963. A survey of Agaricus in Washington, Oregon, and California.
M.S. thesis, University of Washington, Seattle. [unpublished]
Kerrigan RW. 1986. The Agaricales (gilled fungi) of California. 6. Agaricaceae.
Mad River Press, Eureka/ 62 p. + figs.
Kerrigan RW, Callac P, Guinberteau J, Challen MP, Parra LA. 2005. Agaricus
section Xanthodermatei: a phylogenetic reconstruction with commentary
on taxa. Mycologia 97: 1292-1315. http://dx.doi.org/10.1139/b97-058
Norvell L. 2016(1992). The regular column: That nudum is a nuda?
ResearchGate author revision of Mushroom, The Journal. Winter 1991-
92, 10(1): 15-17. <www.researchgate.net/publication/313159493>.
Norvell L. 2017(1988). The herbarium and how it breeds. ResearchGate
author revision of Mushroom, The Journal. Summer 1986, 4(3): 6-9.
<www.researchgate.net/publication/310674716>.
Parra Sanchez LA. 2008. Agaricus L. Allopsalliota Nauta & Bas. Tribu
Agaricaceae S. Imai Parte I. Fungi Europaei 1. Edizioni Candusso.
Parra Sanchez LA. 2013. Agaricus L. Allopsalliota Nauta & Bas. Parte II.
Fungi Europaei 1A. Edizioni Candusso.
Zhao RL, Karunarathna S, Raspé O, Parra LA, Guinberteau J, Moinard M,
De Kesel A, Barroso G, Courtecuisse R, Hyde KD, Guelly AK, Desjardin
DE, Callac P. 2011. Major clades in tropical Agaricus. Fungal Diversity 51:
279-296. http://dx.doi.org/10.1007/s13225-011-0136-7
LORELEI NORVELL
Pacific Northwest Mycology Service
Portland OR 97229-1309, USA
IInorvell@pnw-ms.com
Book Reviews ... 993
Hebeloma (Fr.) P. Kumm. By H.J. Beker, U. Eberhardt & J. Vesterholtt, 2016.
Fungi Europaei 14. Candusso Edizioni s.a.s., <http://eidizionicandusso.it> ISBN
978-88-96059-42-5. 1232 pp, 2654 photos, 200 drawings. 2.8 kilos. Price 86.00 Euros.
Hebeloma is one of those genera in which Tee
species recognition has been difficult, which sericea ete) card
has resulted in a wealth of literature where
authors all interpreted names differently J, )4:15 8e))'//: Vere
and often brought more confusion than
clarification. For Europe, there is now an
excellent book that brings together a mass of
knowledge and data and gives clear reasons for
the species that are presented.
What sets this book apart from other
taxonomic treatments is the care given to
explanations on the decisions made and the
different sets of characters that played a role in
these decisions (biological data concerning monokaryon mating, morphological
data, and molecular DNA data). In some cases morphology tipped the scale
to recognize several species; in others the DNA data indicated the existence
of differences. It should be noted that a combination of molecular markers
was used for species delimitation and that more variable gene regions than
the universally used barcode ITS region were sequenced and phylogenetically
analysed.
The mating data were provided by earlier research (Aanen & Kuyper 1999)
on the Hebeloma crustuliniforme group; the first author contributed most
microscopic observations, whereas the second author brought into the mix
her molecular-phylogenetic skills. Background articles on several sections had
been published previously (e.g. Eberhardt et al. 2013, 2015).
The set-up of the book is comparable to others in the series. The introduction
describes the different chapters and sets the tone. Those chapters cover the
history of the genus, morphology and ontogeny, materials and methods,
infrageneric classification, keys to sections (in different languages), six chapters
treating the species in the various sections, followed by an extensive overview
of ecology and habitat, a list of published names from Europe and north Africa,
bibliography, and finally iconography (more than 500 pages!).
Most important is the chapter called “Our approach’—the materials and
methods part. This is required reading for every taxonomist, and student in
taxonomy. The care with which the research was conducted is exemplary.
994 ... MYCOTAXON 132(4)
Another chapter of enormous importance for taxonomists in and outside
Europe is the list of names that have been published but are not included as
current and valid names in the description chapters. These names come with
annotations of the authors, who studied as many type collections as possible.
The almost 100 pages this list comprises indicates the wealth of information
here (all original diagnoses are given and, if necessary, translated into English).
All 84 species descriptions have been generated in the same way from the
data in the first author's database. This means that they are standardized and
follow the same order.
The keys have also been generated from the database, which means that
human inflation of characters in the keys is absent.
Illustrations in the description chapters are limited to phylogenetic trees
per section or subsection, phenology histograms and distribution maps,
and line drawings of type specimen’s microscopic characters, but the colour
photographs of microscopic and macroscopic characters in chapter 17 add a
wealth of information.
Species recognition still will not be easy in this genus, as the differences
between species can be quite subtle, sometimes only in one morphological
character that needs careful observation, and sometimes there is overlap in
morphological characters among species. This is partly due to the fact that the
authors sought for and found DNA regions that are more variable than ITS.
But remember, this book provides thorough arguments for or against the
decisions made.
It is sad that the third author, Jan Vesterholt, did not live to see the end
product of this work (Jan died in 2011). The book is dedicated to his memory;
Jan brought his knowledge of the northern European species to this project
(Vesterholt 2005).
This book is not the final word on the genus Hebeloma—far from it. The
authors are now focusing on the North American taxa, and I am looking
forward to this next installment. I am very curious to see how many species
the two continents have in common, and whether human introductions might
have played a role in these patterns.
My appreciation and admiration for the authors in tackling this difficult but
important ectomycorrhizal genus hopefully shine through this review.
Aanen D, Kuyper TW, 1999. Intercompatibility tests in the Hebeloma
crustuliniforme complex in northwestern Europe. Mycologia 91: 783-795.
Book Reviews ... 995
Eberhardt U, Beker HJ, Vesterholt J, Dukik K, Walther G, Vila J, Brime
SE 2013. European species of Hebeloma section Theobromina. Fungal
Diversity 58: 103-126. https://doi.org/10.1007/213225-12-0188-3
Eberhardt U, Beker HJ, Vesterholt J. 2015. Decrypting the Hebeloma
crustuliniforme complex: European species of Hebeloma_ section
Denudata subsection Denudata (Agaricales). Persoonia 35: 101-147.
https://doi.org/10.3767/003158515X687704
Vesterholt J. 2005. The genus Hebeloma. Fungi of Northern Europe vol. 3.
Svampetryk, Denmark.
ELSE C. VELLINGA
861 Keeler Avenue, Berkeley, CA 94708 USA
ecvellinga@comcast.net
LICHENS
Lichens of Mexico. The Parmeliaceae - Keys, distribution and specimen
descriptions. Maria Herrera-Campos, Rosa Emilia Pérez-Pérez & Thomas
H. Nash III (eds.). 2016. BristiotHECA LICHENOLOGICA, Vol. 110; vi+723 p.,
17 figs., 14x22cm. English. ISBN 978-3-443-58089-6, bound, price: 199.00 €.
<www.borntraeger-cramer.de/9783443580896> a
Bibliotheca Lichenologica 110
LICHENS OF MEXICO—THE PARMELIACEAE
is a valuable resource that will soon be Fictens oie
consulted even outside of its intended region. ! |
The fact that 20% of the world’s Parmeliaceae
(including the vast majority known from
North America) occur in Mexico make this
volume helpful in keying out lichens outside
of the country, particularly in adjacent Central
America, Caribbean, and United States. Its
use of the recent realignment of species into
phylogenetically supported groups and genera
is a particular benefit.
Thick, dense, but compact, the book is held easily in one hand. Its sturdy
hardcover appears rain- and incidental spillage-proof, and its firmly bound
pages of ‘permanent paper conforming to ISO 9706-1994” do not crackle when
splayed to prepare for heavy use. ‘The text is crisp and legible, although the
spacing is a bit tight for my aging eyes and the margins are far too narrow,
forcing one’s eye to jump occasionally from one long line to another—all
trade offs needed to keep the book affordable and under 800 pages long. This
996 ... MycoTAXxoNn 132(4)
economy of space has the unfortunate result that even when opened firmly,
text on inside margins tends to trail off into the crease. The semi-glossy
photos are sharp and clear but amazingly few: As its title clearly indicates,
the volume focuses on keys and descriptions. Nonetheless, readers who do
read the text should be able to check most of their determinations easily
against the excellent photos in Brodo et al. (2001).
Three introductory chapters set the framework for the 450 species to
follow. The first, covering Mexico’s geology, topography, vegetation, and
diversity, provides four maps, two colored to distinguish the different
climates and biomes. The country’s 2 million km? surface area and varied
landscape, climate, and ecosystems accommodate ~12% of the world’s
species. Herrera-Campos et al. note that it has been historically difficult to
classify Mexico's biological landscape due the convergence of the Holarctic
and Neotropical realms in the Tropic of Cancer, leading to the recognition
of (if not agreement on) 10-32 vegetation types within the five biomes.
The second chapter—Phylogenetic structure of metacommunities in
Mexican Parmeliaceae—presents the first lichen community phylogenetic
analysis for Mexico based on the Parmeliaceae. Liicking et al. note that
the relatively young community phylogenetics field tries to characterize
the phylogenetic structure of ecological communities, thereby ‘providing
insights into evolutionary process’ based on the theory ‘that closely
related species do not usually co-occur in the same community. The few
fungal and micro-organismal studies published before 2006 make these
29 pages the first to apply community phylogenetics to lichens. The 12%-
page table shows the presence/absence of the 450 species in Mexicos five
major biomes—tropical humid forest, humid mountain forest, temperate
forest, seasonally dry forest, and dry shrubland. Species richness and
phylogenetic diversity is highest in the temperate forest and lowest in the
humid mountain and tropical humid forests, supporting the temperate
forest as a center of diversity for Mexican Parmeliaceae while less diverse
and more clade-specific composition of the seasonally dry forest and dry
shrubland favor lineages of the parmotremoid/xanthopermelioid clade
and Oropogon.
The third introductory chapter presents a molecular-based synopsis of
generic classification of Parmeliaceae: since 2000, the ‘increased availability
of DNA sequence data has led to a second revolution of generic delimitations
in Parmeliaceae based on molecular phylogenetic studies addressing the
circumscription of genera’ Crespo et al. briefly evaluate currently accepted
Book Reviews ... 997
genera, recapitulating recent changes in their circumscription and dividing
the family into 7 phylogenetic groups:
ALECTORIOID—Circumscriptions of Alectoria, Bryoria, and Pseudephebe
remain relatively unchanged. The 2009 separation of Gowardia from Alectoria
is not currently supported, while Nodobryoria, segregated previously from
Bryoria, appears not closely related to Bryoria.
CeTRARIOID—Cetraria, Kaernefeltia, | Melanelia, Tuckermanella,
Tuckermannopsis* [sic], Vulpicida represent a usually well-supported
monophyletic clade, although current circumscriptions are needed in some
instances. “Genetic distances among many currently accepted genera are
remarkably low.... Thus at least some of the genera should most likely be
merged.” In this chapter Crespo & Barrano propose the new Kaernefeltia
iberica comb. nov. for extralimital Spanish specimens previously referred to
K. merrillii, a North American endemic.
Hypocymnioip— The first sentence reads: “this group consists of four
foliose genera’, but only three are named (probably because the authors counted
Cavernula, whose species were transferred to Hypogymnia in 2011). Brodoa
[not found in Mexico] and Pseudevernia are the other two hypogymnioid
genera cited here.
LETHARIOID—Segregation of Letharia (found in Mexico) and Lethariella
awaits further research.
PARMELIOID— This largest group accommodates over 1800 accepted
species. Of the 21 accepted genera, 5 (Bulbothrix, Parmelia, Parmelinella,
*NOMENCLATURAL NOTE: Although the genus is misspelled ‘Tuckermannopsis’
consistently throughout the volume, according to Art. 60.1 of the INTERNATIONAL
CODE OF NOMENCLATURE FOR ALGAE, FUNGI, AND PLANTS (McNeill & al. 2012),
the name honoring Edward Tuckerman should be corrected to Tuckermanopsis, as
originally spelled for the type species Tuckermanopsis ciliaris (Ach.) Gyel. and as
displayed on Index Fungorum <http://www.indexfungorum.org> and MycoBank
<www.mycobank.org>. MycoBank lists “Tuckermannopsis’ as an orthographic variant.
Thus far I have been unable to determine just exactly when the name acquired
the extra ‘n’ but suspect it might have been introduced by Hale, who transferred 13
species to the genus in the 5" lichen checklist by Egan (1987). Although a Google search
tends to turn up more Tuckermannopsis hits, the spelling used almost uniformly in the
North American books on my library shelves (e.g., Hale & Cole 1988, Brodo et al. 2001
McCune & Geiser 2009), there are quite a few papers (e.g., Harris & Lendemer 2005;
Randlane & Saag 2006 + many more European texts) that spell the name as proposed
by the Byelorussian Gyelnik in 1933. Note also that when Esslinger (2006) carved the
segregate genus, Tuckermanella Essl. from Tuckermanopsis, he correctly spelled his new
genus with only one ‘n.
998 ... MycoTAXxoON 132(4)
Parmotremopsis, Pseudoparmelia) are currently polyphyletic, in a state of flux,
or lack molecular data; three (Canoparmelia, Hypotrachyna, Xanthoparmelia)
were recently trimmed for monophyly, and three (Melanelixia, Melanohalea,
Remotrachyna) are recent segregates. The remaining 10 well-supported
monophyletic genera include Cetrelia, Flavoparmelia, Montanelia, Myelochroa,
Parmelina, Parmeliopsis, Parmotrema, Phacopsis, Punctelia, and Relicina.
PROTOPARMELIOID—confined to the polyphyletic Protoparmelia.
UsnEoip—limited to one genus (Usnea) with 350 species and the most
highly diverse in tropical areas. Usnea now includes previous segregates
(e.g., Dolichousnea, Eumitria, Neuropogon) phylogenetically nested within it.
GENERA WITH UNCERTAIN AFFINITIES—Anzia, characterised by multispored
asci with small curved ascospores and a spongiostratum, is centered in eastern
Asia. Imshaugia is distinguished by Parmeliopsis-like species with emergent
laminal/marginal pycnidia and bacilliform/bifusiform conidia. Menegazzia,
diagnosed by a perforated upper thallus surface, occurs primarily in the
southern hemisphere. The morphological segregation of Nodobryoria from
Bryoria is molecularly supported. Oropogon a fruticose lichen of the Neotropics
with a currently unresolved phylogeny.
A key to the identification of genera representing Parmeliaceae in Mexico
concludes the introduction.
A formidable array of Parmeliaceae experts have written chapters for this
volume, with Editors Herrera-Campos (5), Pérez-Pérez (9), and Nash (24)
and authors R.S. Egan (5), T.L. Esslinger (10), J.A. Elix (4) each contributing
to more than two chapters while J.W. Bjerke, A. Crespo, P.K. Divakar, S.D.
Leavitt, James Lendemer, Robert Liicking, H.T. Lumbsch, Bruce McCune,
I.KK. Tronstad, J.L. Vilmasenor, and A. Zambrano Garcia each contributed
to at least one.
Thirty-nine generic chapters covering 450 species comprise the bulk (and
the goal) of THE PARMELIACEAE. The note added in proof at the bottom of
the section’s title page hints at editorial headaches suffered over collating
over 40 papers, with many undoubtedly arriving only shortly before press
time:
“Herein Dr. Egan proposes treating everniastrum subplanum
Sipman as a synonym of Parmotrema paramoreliense whereas
the former species was treated as Hypotrachyna subplanum in the
Hypotrachyna treatment as proposed by Divakar et al. (2013a). The
correct placement must await molecular investigations of all relevant
material”
Book Reviews ... 999
Genera are presented in alphabetical order and entitled with the genus name
+ in Mexico (except for “Menegazzia in Mexico and the Caribbean islands”).
The chapters range in length from one page (Cetraria, Phacopsis) to 103 pages
(Hypotrachyna, Parmotrema), depending on the number of species. Formatting
is consistent: Title and author information are followed by abstract, short
genus characterization, and species key (for genera with more than one
species covered). Each species treatment begins with the taxonomic
heading and nomenclator followed by morphological description, chemical
composition, distribution data, and important notes and concludes with
a list (often extensive) of specimens examined. Alphabetical presentation
—a necessity in such a large volume with no index—and running titles
enable rapid location of a particular genus. Eight species new to science
are proposed in the Alectoria (2), Hypotrachyna (1), Parmotrema (2),
Pseudevernia (2), and Tuckermanella (1) chapters.
There are a few deviations, perhaps the most startling being that only the
species key, taxonomic headings, illustration references, and distributional
data appear are provided for Canoparmelia; I regret the absence of the rest
of the material but suspect that the fact this chapter was written by two
already overcommitted editors, Pérez-Pérez & Nash, may have something to
do with the decision to withhold the morphological and chemical features
[referenced in the abstract as “Descriptions were provided previously”] and
final discussions for the 13 species. Still, it will be a bit of a hassle for some
readers to find the correct ‘previous’ paper for the missing information.
I also greatly mourn the lack of a concluding comprehensive species
index that would allow easy tracking of new taxonomic realignments,
always helpful to those wondering what name a favored species has recently
adopted. The information can be ferreted out of the nomenclators, although
the reader must know the new name to find out what happened to the old
one. This does, however, have the benefit of persuading the user to consult
the keys, the whole purpose of the volume!
Finally, given the focus on phylogenetic analysis of biomes, groups,
and genera in this volume, I was exasperated by the lack of information
regarding which DNA regions were sequenced to generate the phylogenies.
A time-consuming scan of the 31-page bibliography proved somewhat
helpful, although I found only three titles naming the DNA regions:
mitochondrial DNA (Crespo & al. 2001), ITS & {-tubulin (Crespo & al.
2002), and ribosomal markers + RPB1 (Crespo & al. 2007). Inspection of
the multilocus sequences analyzed by Altermann et al. (2014), however,
1000 ... Mycotaxon 132(4)
explains why modern titles do not enumerate all DNA regions used: citing
their 15 different loci (even in the abstract) would have needlessly lengthened
the paper and the authors cited the needed information in their Materials
& methods table. Nonetheless, the nuclear ITS, LSU and mitochondrial
SSU rDNA sequences cited in the abstract for “DNA barcoding in the
Hypotrachyna clade” abstract (Divakar et al. 2015) is information that
should have been shared under Hypotrachyna in the third chapter. Likewise
identifying the DNA barcode used to identify hidden diversity in Parmelia
sensu stricto (Divakar et al. 2015) as derived from the ITS region should have
been noted for that genus.
The rapid change in the complex molecular landscape is perhaps one
reason why most authors use vague terms such as ‘molecular analyses,
‘DNA sequence based, or ‘phylogenetic analyses’ in their titles (saving the
lengthier specifics for the abstracts). If a second edition is anticipated, a
brief list of the DNA regions sequenced used to generate the phylogenetic
support and their utility (even if just in table form) would be welcome.
The objections raised above are minor and should in no way discourage
anyone from purchasing such a useful book that does precisely what it
sets out to do: key and describe 450 lichens representing Parmeliaceae
from Mexico. As noted above, its use will not (nor should) be restricted to
Mexico and the reference will prove a reliable resource for all those who use
it. 1am very happy to have LICHENS OF MExIco: THE PARMELIACEAE On my
lab library shelves.
Altermann S, Leavitt SD, Goward T, Nelsen MP, Lumbsch HT. 2014. How
do you solve a problem like Letharia? A new look at cryptic species
in lichen-forming fungi using Bayesian clustering and SNPs from
multilocus sequence data. PLoS One 9: e97556.
https://doi.org/10.1371/journal.pone.0097556
Brodo IW, Sharnoff, SD, Sharnoff S. 2001. Lichens of North America. Yale
University Press, New Haven. 795 p.
Crespo A, Blanco O, Hawksworth DL. 2001. The potential of
mitochondrial DNA for establishing phylogeny and _ stabilising
generic concepts in the parmelioid lichens. Taxon 50: 807-819.
https://doi.org/10.2307/1223708
Crespo A, Molino MC, Blanco O, Schroeter B, Sancho LG,
Hawksworth DL. 2002. rDNA ITS and f-tubulin gene sequence
analyses reveal two monophyletic groups within the cosmopolitan
lichen Parmelia_ saxatilis. Mycological Research 106: 788-795.
https://doi.org/10.1017/S095375620200610X
Book Reviews ... LOO1
Crespo A, Lumbsch HT, Mattson JE, Blanco O, Divakar PK, Articus
K, Wiklund E, Bawingan PA, Wedin M. 2007. Testing morphology-
based hypotheses of phylogenetic relations in Parmeliaceae
(Ascomycota) using three ribosomal markers and the nuclear
RPB1 gene. Molecular Phylogenetics and Evolution 44: 812-824.
https://doi.org/10.1016/j.ympev.2006.11.029
Divakar PK, Leavitt SD, Molina MC, Del-Prado R, Lumbsch HT, Crespo
A. 2015. A DNA barcoding approach for identification of hidden
diversity in Parmeliaceae (Ascomycota): Parmelia sensu stricto as
a case study. Botanical Journal of the Linnean Society 180: 21-29.
https://doi.org/10.1111/boj.12358
Divakar PK, Crespo A, Nufiez-Zapata J, Flakus A, Sipman HJM, Elix J,
Lumbsch HT. 2013. A molecular perspective on generic concepts in the
Hypotrachyna clade (Parmeliaceae, Ascomycota). Phytotaxa 132: 21-38.
https://doi.org/10.11646/phytotaxa.132.1.2
Esslinger TL. 2003. Tuckermanella, a new cetrarioid genus in western North
America. Mycotaxon 65: 135-141.
Hale M. 1987. 13 Tuckermannopsis [sic] recombinations proposed on p. 165
in Egan RS. A fifth checklist of the lichen-forming, lichenicolous and
allied fungi of the continental United States and Canada. The Bryologist
90(2): 77-173.
Harris RC, Lendemer JA. 2005. Contributions to the lichen flora of
Pennsylvania: A checklist of lichens collected during the first Howard
Crum Bryological Workshop, Delaware Water Gap National Recreation
Area. Opuscula Philolichenum 2: 1-10.
McCune B, Geiser L. 2009. Macrolichens of the Pacific Northwest. Revised
Edition. Oregon State University Press, Corvallis.
McNeill J, Barrie FE, Buck WR, Demoulin V, Greuter W, et al. (eds).
2012. International Code of Nomenclature for algae, fungi, and plants
(Melbourne Code). [Regnum Vegetabile no. 154.] Koeltz Scientific
Books, Konigstein.
Randlane T, Saag A. 2006. Cetrarioid lichens i34n Europe - an identification
key for the species. 75-84 in: Lackovicova A., Guttova A., Lisicka E.
& Lizon P. (eds.), Central European lichens - diversity and threat.
Mycotaxon Ltd., Ithaca.
LORELEI NORVELL
Pacific Northwest Mycology Service
Portland OR 97229-1309, USA
IInorvell@pnw-ms.com
Russula vinosoflavescens sp. nov.
(Trendel, Hampe & Verbeken— Fie. 1, p. 713)
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